BLUMEA 30 (1985) 279-318

The Malesien species of Paspalum L. (Gramineae)

R. de Koning & M.S.M. Sosef

Rijksherbarium, Leiden, The Netherlands

Summary

Paspalum L. (Gramineae: Paniceae) has 14 species in Malesia, 5 of which are cultivated only.

In the P. scrobiculatum-complex 5 varieties have been distinguished. One of these (from Africa)

needed a new name: var. lanceolatum Koning & Sosef, another (from Southeast Asia) a new

combination: var. horneri (Henr.) Koning & Sosef.

Introduction

Paspalum was described by Linne (1759), who included four species in it: P. dimi-

diatum, P. distichum, P. paniculatum, and P. virgatum. The first name was a new one

for Panicum dissectum L. and therefore superfluous, an error he himself corrected in

1762. first As this was the species described by him, it has generally been accepted

of the The of as the type generic name. use P. virgatum (e.g. Tsvelev, 1976) is there-

fore incorrect.

authors have the Subsequent substantially expanded genus, partly by describing

new species, partly by the inclusion of species first included in genera as Axonopus

Beauv., Cynodon Pers., Digitaria Haller, Eriochloa H.B.K., and Syntherisma Walt.,

whereby at present there are 200 (Jacques-Felix, 1962) to 250 (Clayton & Renvoize,

1982) species, most of them American. Unfortunately since Flugge's revision (1810)

no attempt at an overall study has been madeexcept for compilatory works as those

by Roemer & Schultes (1817), Kunth (1833), and Steudel (1853). Most other work

has been done on a regional scale as is the case in the present one. A major review is

that of Chase (1929) for the North American taxa which, because of its thorough-

ness, must serve as the base for all subsequent work. Most South American taxa have

African been treated by Parodi (1937). The tropical ones have been revised by Clay-

the the ton & Renvoize (1982), Indian ones by Bor (1960), and Australian ones by

Vickery (1961, 1975). Especially the latter four publications were ofimportance, as

they included most of the species also occurring in Malesia.

INTERGENERIC RELATIONSHIP

Paspalum is a member of the Paniceae and is usually regarded to be closely related to Panicum L. The principal generic character at first used by Linne to distinguish 280 BLUMEA - VOL. 30, No. 2, 1985

the two was the absence of the lower glume in Paspalum. As Chase (1911) has shown

feature unreliable numberof included this is since in quite a taxa presently in Paspa-

lum this scale is more or less developed, either occasionally so (P. distichum in Male-

sia), or as a rule (P. decumbens Sw.).

Since Linne's time Paspalum has generally been regarded as a genus in its own

rights. According to Chase (1911; 1929) it is distinct by the strictly racemose inflo-

the rescences, plano-convex or slightly concavo-convex spikelets, which are abaxial,

with the lower sterile lemma from fertile i.e. glume and turned away the rachis, the

lemma with involute, not infolded or straight margins, and the obtuse, indurated fer-

tile floret in fruit When this there have been few if (fig. 5c). diagnosis is applied any

whether problems to recognize a species belongs to Paspalum or not.

INFRAGENERIC DELIMITATION

Chase has divided the North American number of (1929) species into a groups,

which system was more or less copied by Pilger (1940) in his global survey of the

first the of of genus. At sight presence solitary or geminate spikelets, or glabrous or

hairy ones would suggest some sort of affinity. We think, however, that Chase was

when she these characters her correct nearly completely neglected in arrangement.

For example P. orbiculare Forst.f. has both solitary and geminate spikelets within

the same raceme, while P. longifolium Roxb. may have strongly pubescent to less

pubescent, rarely even entirely glabrous spikelets. The two seem to us to be closely

related because of the shape of the spikelets and the three distinctly dark-coloured

of the and sterile lemma. As have dealt here with few nerves upper glume we only a

taxa, no opinion on their distinctiveness is expressed. To exclude any suggestion of

relationship we have arranged our taxa alphabetically.

Tsvelev (1973, 1976) has distinguished next to the typical section a section Diplo-

stachys (Steud.) Tsvelev for 'all species with two digitately arranged racemes', typi-

fied by P. distichum. Such a distinction may be satisfactory for the U.S.S.R. with

only four species, but on a worldwide scale the only distinction given is insufficient.

SPECIFIC AND INFRASPECIFIC DELIMITATION

The Malesian taxa in general have offered little problems in their delimitation,

of for the - except course notorious P. scrobiculatum complex.

This is a very variable group both in Malesia and elsewhere. Several authors have

of ranks within but all failed reach distinguished taxa various it, to a satisfactory result. Henrard stated that 'this of the of (1940) aptly is one most puzzling groups

do dare claim that the solution grasses.' Similarly we not to adopted here can be the

final one although we of course think that it is the most acceptable and practical one.

The names usually found in literature of the taxa involved are P. auriculatum

Presl, P. cartilagineum Presl, P. commersonii Lamk., and P. scrobiculatum L. If only

the type material of these names is inspected it is easy to believe that they indeed R. de Koning & M. S. M. Sosef: Malesian species of Paspalum 281

represent different species. Additional material, however, brings them closer and

closer and usually a complete range from one extreme to the other can be laid out

in the end. To us it now seems that we have been dealing with a single species dis-

playing an enormous variation. Singh's (1965) suggestion for African representatives

further that they represent an apomictic swarm deserves research. The situation ob-

served in the Bothriochloa bladhii-complex (De Wet & Harlan, 1970, as B. inter-

media) is brought to mind, although that may even be more complicated, as what

many have considered to be generic boundaries, are transgressed there.

To reduce all to a single species without further ado seemed to be a rather easy

mathematical way out, especially when after the classical, descriptive approach a

elaboration of the material which showed the of number of was made, presence a

clusters in which about all of the specimens could be accommodated. Considering

the final result we fully agree with Hooker f. (1896) who stated in another context

that 'no two botanists working independently over the same material, would arrive

at the same (conclusion), or agree in any other.'

Of the available for of numerical many possible analyses studying homogeneity

modified for data we chose Principal Component Analysis (Sneath & Sokal, 1979),

computer analysis by Zandee (cf. Zandee & Glas, 1982), whereby the possible exis-

of clusters detected. The considered tence can relatively easily be specimens are to rest in where the of to the a multidimensional space number dimensions is equal

number of characters. The directions in which the largest, the one but largest, etc.

variances occur can then be determined. These directions are called the 'principal axes' (P. A.). The projection of the specimens on these axes are called 'principal

components' (P.C.). A large variance occurs when many specimens are far off the

i.e. in mean, where clustering takes place (roughly comparable to solar systems

stellar The ones in galaxies in super-galaxies). analysis shows how much each charac-

ter contributes to each principal axis, the so-called 'variation load', i.e., the degree of

ifcharacter 'x'has load of variability present. For example, a variation 80% in P. A. 'a',

this that of the of'x' P. visual- means 80% variation is represented in A. 'a', or, more

ly, that P. A. 'a' lies in about the same direction as the axis in the multidimensional

space along which 'x' is represented. The higher the variation load, the more impor-

tant that character is in that P. A. In this way 'heavy' characters are suggested. How-

ever, it must be realized that the load is composed of two parts: the variation within

a cluster (internal variation) and ofthat between clusters (external variation). When a

taxon single is very variable for a certain character a large variation load will result, but it is internal and for the delimitationof taxa it is then useless. The ones with the

largest external variation which will cause disjunction between clusters can be detect- ed by drawing plots in which the direction of each character (the direction cosine) is

set out. On each P.A. each character has its own 'Eigenvector', a transformation of its variation load. In diagram 1 the P.A.'s are divided into arbitrary units to accom-

modate these values, from which then perpendiculars are drawn. From the point

(0.0) derived vectors can be drawnto the resulting coordinates. These last vectors are measures of the external variation of the characters. Those that are both long and 282 BLUMEA - VOL. 30, No. 2, 1985

Table 1. Characters used for the numerical analysis. For the dimensions the unit of

approximation is given between brackets.

1. Plant length (cm)

of sheath of the third leaffrom 2. Length the top (cm)

of 3. The pubescence 2: 1 = glabrous

= at 2 margin hairy the apex

3 = margin hairy all along

4 = hairy all over

4. Length of its ligule (0.1 mm)

5. Length of its blade (cm)

6. Ratio oflength of blade/sheath

7. Width of this blade (mm)

8. Length/width ratio of blade

of blade: short hairs behind the 9. Pubescence 1 = ligule

2 = long hairs behind the ligule

3 = as 2, lower part margin hairy

4 = idem, lower part and midrib hairy

5 = blade hairy all over

10. Diameter peduncle, 1 cm from top (0.1 mm)

11. Number of racemes/inflorescences

12. Length of lowest raceme (mm)

13. Width of rachis (0.1 mm)

14. Length of spikelet (from middle of raceme) (0.05 mm)

15. Width of spikelet (idem)

16. Length/width ratio of spikelet

Number of of the 17. nerves upper glume

18. Number of nerves of the sterile lemma

19. Texture sterile lemma: 1 = membranous

2 = slightly indurated

3 = indurated

of this from point closest to the centre a cluster separate others. The more the vec-

towards the the the correlation tors point same direction, greater positive between the characters they represent.

of hundred A random sample a specimens sufficiently representing P. auricula- turn ‘P. commersonif and ‘P. scrobiculatum’ was drawn whereby only complete material was included. A number of characters were measured, standardized and checked for their variation loads in the first five P.C.'s. The nineteen most important ones used for the preparation of the final diagrams are listed in table 1. R. de Koning & M. S. M. Sosef: Malesian species of Paspalum 283

Table 2. Percentage variation load of original characters in the first principal

components.

Character P. A. 1 P. A. 2 P. A. 3 P. A. 4 PA. 5

1. 66.72 3.42 0.07 3.46 2.98

2. 61.06 5.51 8.81 9.06 2.36

3. 25.69 19.38 0.31 8.74 13.68

4. 19.74 3.21 9.35 1.43 33.25

5. 81.04 0.01 3.27 5.35 0.95

6. 8.76 1.83 0.42 66.08 0.11

7. 68.42 4.90 13.09 3.74 0.05

8. 8.86 12.10 50.92 0.00 3.21

9. 28.24 7.51 4.24 4.25 23.45

10. 67.97 3.82 6.98 0.94 0.50

11. 48.72 9.30 15.20 0.65 0.00

12. 75.47 2.32 0.00 0.04 2.17

13. 38.18 15.36 2.80 3.18 0.02

14. 0.36 77.08 0.09 0.90 3.27

15. 1.45 78.72 9.07 0.38 0.24

16. 1.81 8.13 33.74 0.24 21.22

17. 0.67 58.38 0.54 5.84 1.01

18. 1.31 46.12 1.04 5.94 0.34

19. 1.35 4.57 19.84 14.27 10.27

Table 2 the variation loads for these features. The the presents largest variance in

data, shown by the first P. A., is to be found especially in the length and width of the

leaf, diameter of the peduncle (an unusual feature), and in the length of the raceme.

The second P. A. contains variation of the spikelet characters (dimensions, number of nerves). The third P.A. derives its importance mainly from the shapes of the

leaves and of the spikelets.

Since each specimen occupies its own place along each P.A. it is possible to visual-

ize the clusters possibly present by plotting pairs of P.A.'s against each other. Dia-

gram 1 shows three such plots, i.e., those of the first three P.A.'s.

Table 3 presents the standardized, normalized Eigenvectors for the first five P.A.'s

from which the combined vectors in the plots have been derived.

In the first plot (diagram la, P.A. 1/P.A. 2) three clusters can be distinguished, one

containing all ‘P. scrobiculatum’ specimens, another those of ‘P. commersonii’and a

' third those of ‘Pauriculatum’. In the last one the grouping around a distinct main

centre is weak and one might still doubt whether it really is disjunct from the ‘P. commersonii ’-cluster. - 284 BLUMEA VOL. 30, No. 2, 1985

Table 3. Standardizednormalized Eigenvectors.

Character P. A. 1 P. A. 2 P. A. 3 P. A. 4 P. A. 5

1. 0.9073 -0.2083 -0.0374 0.2289 0.2996

2. 0.8680 -0.2647 0.4159 0.3703 0.2662

3. -0.5630 0.4962 -0.0777 -0.3637 0.6413

4. 0.4935 0.2019 0.4285 -0.1469 -1.0000

5. 1.0000 -0.0089 0.2536 -0.2846 0.1688

6. 0.3288 0.1523 0.0911 -1.0000 -0.0577

7. 0.9188 0.2495 -0.5070 -0.2379 0.0381

8. 0.3307 -0.3921 1.0000 0.0051 0.3109

9. -0.5903 0.3088 -0.2886 -0.2537 0.8397

10. 0.9158 0.2204 -0.3704 -0.1196 0.1228

11. 0.7754 -0.3437 -0.5464 -0.0995 0.0117

12. 0.9650 0.1717 -0.0094 -0.0255 0.2557

13. 0.6864 0.4417 -0.2344 0.2193 -0.0271

14. 0.0666 0.9895 0.0414 0.1165 -0.3135

15. 0.1338 1.0000 0.4220 0.0754 0.0846

16. -0.1496 -0.3214 -0.8140 0.0607 -0.7989

17. -0.0906 0.8612 -0.1029 0.2974 -0.1739

18. 0.1271 0.7654 -0.1426 0.2998 0.1013

19. 0.1289 -0.2409 -0.6241 0.4647 0.5558

The second plot (diagram lb, P. A. 1/P.A. 3) shows no really disjunct clusters.

The third plot (diagram lc, P.A. 2/P.A. 3), however, shows two more obviously

ones. One contains all of ‘P. auriculatum’ and ‘P. scrobiculatum disjunct specimens ’,

the other one all those of ‘P. commersonif’.

This separation is especially due to the dimensions of the spikelets (characters 14

of the husks The and 15) and the number nerves in (characters 17 and 18). apparent

between the two clusters lies at 2.6 2.15 mm. After intensive gap spikelet-size by study of all the material, both that used for the analysis and that of the remainder, it

turned out that few with this set of dimensionscould only very specimens particular be found. There was more overlap in the number of nerves (5—7 versus 7—13).

which In this way it has become clear that there are three distinct clusters present

three-dimensional with the three classi- might be depicted in a plot. They correspond

cal taxa.

Characters to distinguish between them are in decreasing importance of their con-

tribution to the first three P.A.'s: width of the spikelet (character 15), widthof the blade (7), its length (5), diameter of the peduncle (10), length of the lowermost raceme (12), length of the spikelet (14), length of the third sheet from top (2), num- R. de Koning & M. S. M. Sosef: Malesian species of Paspalum 285

Diagram 1. Paspalum scrobiculatum L. Plots of the first three principal componentsagainst each

other. ■ var. auriculatum (‘P. auriculaturn’), • var. bispicatum (‘P. commersonii’),* var.

scrobiculatum (‘P. scrobiculatu’).

these to ber of racemes (11), etc. Since do not appear be very impressive features it

seemed best to regard the clusters as varieties. These are then to be called P. scrobicu-

latum var. scrobiculatum, var. auriculatum (Presl) Merr. and var. bispicatum Hackel.

The latter is the required combination for ‘P. commersoni’ as P. scrobiculatum var.

commersonii (Lamk.) Stapf (1920) is antedated by Hackel's combination of 1914

proposed for specimens that are fairly aberrant by the presence of two subopposite racemes, but otherwise belong to this taxon. The well-known epithet ‘commersonii’

can therefore unfortunately not be retained.

We have mentioned the of a called ‘P. cartilagineum which presence species ’, sup- of posedly would be distinct by the presence an indurated sterile lemma. The speci- 286 BLUMEA - VOL. 30, No. 2, 1985

mens thus identified turned out to be spread out rather evenly over the three other

itself cluster of varieties, the type lodging in the var. bispicatum. Moreover, some

with non-induratedlemmas within the specimens have been seen induratedand same

inflorescence whereby the value of this character is brought to nought and cannot be

maintainedto distinguish a taxon.

The large variation found within this complex in Malesia may be caused partly by

field ecological effects, e.g., soil type, shade, moisture, etc. A study wouldresult in a better understanding of the causes and a possible explanation for the intermediary

forms occasionally encountered.

that similar the continental It is interesting to note a variability is present in Asian

material, but that in Africa the situation seems to be different (as far as could be judged from literature and the material available in L). Here var. bispicatum and an- other taxon of the complex, which is not represented in Asia, occur. This latter form

has unfortunately been confused with P. auriculatum (Clayton, 1974; Singh, 1965;

Clayton & Renvoize, 1982), but is quite different from that because of the rather loose sheaths, the relatively short and very wide leaves, with a rounded base, and

spikelets up to 2.5 mm long. These characters are similar to those used to distinguish

the varieties in P. scrobiculatum and we therefore regard this taxon not as a species

has been done the authors mentioned but also as by above, as a variety. A correct

combination seems to be lacking for it and we here therefore propose the combina-

tion P. scrobiculatum var. laneeolaturn Koning & Sosef (see Appendix, p. 312, and

the Key, p. 288).

ALIEN AND EXCLUDED SPECIES

A number of species occur only as cultigens apparently without escaping. They

are: P. decumbens Sw., P. notatum Flugge, P. plicatulum Michx., P. virgatum L., and

P. wettsteinii Hackel. These have not been treated here formally but to avoid confu-

identification with the also sion in the native or escaped taxa (and to hedge our bets

included them in case they may escape in the future) we have in our key and in some

notes at the end of this paper.

CHROMOSOME NUMBERS

We have not made any counts ourselves, but have cited those reported in the liter-

The basic be for ature. number apparently is x = 10, as was to expected a panicoid

P. orbiculare, genus. Aneuploidy (2n = 54) has been observed in q.v. Unfortunately

no vouchers have been seen. In some cases misidentifications seem unlikely (e.g. of

P. dilatatum and P. while others P. conjugatum, paniculatum ), in descriptions were

provided that showed how the name used was to be applied. Within the P. scrobicula-

tum -complex however, there is less certainty. In the case of var. bispicatum one can , usually be fairly sure that the material was correctly identified, but for the other ones this is doubtful and the data for the latter have therefore been omitted. Counts of 60 have been made for the Hsu 2n = 40, aggregate species, e.g. by (1971). R. de Koning & M. S. M. Sosef: Malesian species of Paspalum 287

ACKNOWLEDGEMENTS

carried the ofthe for The present study was out at Rijksherbarium(L) as part requirements a

M. Sc. degree at the Rijksuniversiteit, Leiden, the Netherlands, under the supervision of Dr. J.F.

Veldkamp. We are very grateful for the latter's advice and his extensive polishing of the final

manuscript. We would like to thank Dr. M. Zandee and Dr. D. Povel, Leiden, for their helpful

comments and critical discussions on the intricacies of numerical taxonomy. The research was

We of the Rijks- primarily based on material present in L. are therefore most obligedto the Director

herbarium and his staff for the opportunities given and also to the Keepers and Directors of

B, BM, BO, G, K, MICH, P, PR, SING, UPS, US, and W, who kindly lent additional collections

which have made in Dr. L.E. unable without the study could not been a satisfactory way. Kers, STB,

to send the type material ofPaspalum conjugatum (q.v.) kindly had photocopies made, which

proved to be quite sufficient to identify the name. Negatives of the specimens in the Lamarck her-

trans- barium, Paris, were also gratefully received. Dr. J.van Brummelen,Leiden, kindly provided

lations of Russian text. The drawings and diagrams presented here were prepared by ourselves.

REFERENCES

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1911. Notes of Paniceae. IV. Proc. Biol. Soc. 24:137-141. CHASE, A. on genera Repr. Washington

1929. The North American species of Paspalum. Contr. U.S. Nat. Herb. 28 (1): xvii +

310 pp.

CLAYTON, W.D. 1974. The Paspalum scrobiculatum complex in tropical Africa. Kew Bull. 30:

101-105.

3:607-612. — & S.A. RENVOIZE. 1982. Flora of tropical East Africa, ed. 2,Gramineae London.

FLUGGE, D.J. 1810. Graminum monographiae 1, Paspalus, Reimaria. 244 pp. Hamburg.

3: 411-480. HENRARD, J.Th. 1940. Notes on the nomenclature of some grasses. Blumea

HOOKER, J.D. 1896. Flora of British India 7: 14. London.

and HSU, C. 1971. A guide to the Taiwan grasses, with keys to subfamilies, tribes, genera spe-

cies. Taiwania 16: 290-294.

JACQUES-FELIX, H. 1962. Les Graminees d'Afrique tropicale 1: 239-240. Paris.

KUNTH, C.S. 1833. Enumeration plantarum 1: 40-65. Stuttgart.

LINNE, C. 1759. Systema Naturae, ed. 10, 2: 855. Stockholm.

1762. Species plantarum, ed. 2: 81. Stockholm.

1937. Contribucion al estudio de las Gramineas del de la flora PARODI, L. R. genera 'Paspalum'

Uruguaya. Rev. Mus. La Plata n.s. 1: 211-250.

PILGER, R. 1940. Gramineae III, Unterfamilie Panicoideae. In: A. Engler & K. Prantl, Die

Natiirlichen Pflanzenfamilien ed. 2,14e: 58-67. Leipzig.

ROEMER, J.J. & J.A. SCHULTES. 1817. Systema vegetabile 2: 290-317. Stuttgart.

SINGH, D.N. 1965. Apomixis in Paspalum II: Paspalum commersonii Lamk. Rev. Biol. Lisboa 5:

75-83.

1979. Numerical + 573 San Francisco. SNEATH, P.H.H. & R.R. SOKAL. taxonomy, xv pp.

STEUDEL, E.G. 1853. Synopsis plantarum glumacearum 1: 16-33. Stuttgart.

TSVELEV, N.N. 1973. Notae de gramineis florae URSS, 7. Nov. Syst. PI. Vase. 10: 79-80.

1976. Grasses of the Soviet Union: 667-669. Leningrad. (English translation, 1983: 1006-

1009, New Delhi.)

VICKERY, J.W. 1961. Gramineae. Contr. New South Wales Nat. Herb., Fl. Ser. 19: 115-122.

1975. Gramineae. Fl. New South Wales no. 19: 135-136. Sydney.

WET, J.M.J. DE & J. R. HARLAN. 1970. Bothriochloa intermedia. A taxonomic dilemma. Taxon

19: 339-340.

ZANDEE, M. & P.C.G. GLAS. 1982. Studies in the Holcus lanatus-Holcus mollis complex (Poa-

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Wet. C, 85: 413-437. 288 BLUMEA - VOL. 30, No. 2, 1985

PASPALUM

Paspalum L., Syst. Nat. ed. 10, 2 (1759) 855; Flugge, Gram. Monogr., Paspalum (1810) 53;

R. & S., Syst. Veg. 2 (1817) 13; Nees, Agrost. Bras. (1829) 18; Steudel, Syn. 1 (1853) 16;

Benth. & Hook.f., Gen. PI. 3 (1883) 1097;Chase, Proc. Biol. Soc. Washington 24 (1911) 137;

Contr. U.S. Nat. Herb. 28, 1 (1929) 7; Parodi, Rev. Mus. La Plata n.s. 1 (1937) 211; Pilger in

E. & P., Nat. Pfl. Fam. ed. 2, 14e (1940) 58. — Sabsab Adans., Fam. PI. 2 (1763) 31, 599,

= nom. superfl. - Lectotype: P. dimidiatum L., nom. superfl. P. dissectum L. (Typotype:

Pluk., Almagesti... Mantissa (1700) 94, t. 350, f. 2.)

For further Chase synonyms see (1929).

The following description is based on the Malesian taxa only. Some of the specific

characters they have in common have been included here to prevent unnecessary repetitions.

Annuals or perennials, branching intra-vaginally at base. Sheaths slightly compres-

sed. Ligule entire, membranous, glabrous. Blades erecto-patent, margins scabrous.

Peduncle exserted, smooth. Racemes 1-many, spreading to drooping at maturity, all

shortly stalked. Stalk terete. Rachis subterete to distinctly winged. Spikelets abaxial,

plano-convex or sometimes unequally biconvex or concavo-convex, subsessile or

shortly pedicelled, falling as a whole, solitary or paired (once ternate) (hence 'inner' and 'outer' spikelets), in two rows on one side of the rachis, the groups alternate. Ar-

ticulation of pedicel straight, top broadened. Lower glume usually absent. Upper glume present, membranous, smooth, margins usually involute, rarely flat, outer

Sterile lemma the nerve (s) submarginal. as upper glume, epaleate. Fertile lemma

shorter than the slightly spikelet, chartaceously indurated, rarely but slightly so, margins involute, germination flap long-trapezoid, c. 0.8 times as long as the lemma, its margins usually visible as pale lines. Palea similar to and enclosed by the lemma

shorter, involute at the and the lower (fig. 5c) slightly margins membranous, apex in half. Lodicules obtrapezoid, thin, veinless, free, glabrous. Anthers 3. Styles 2, free at base. Embryo c. 0.33 times as long as the caryopsis. Hilum subbasal, oblong.

Distribution. About 200—250 species, mainly tropical, 14 in Malesia, of which

5 apparently native or archaeophytical, 4 introduced and escaping, 5 only known from cultivation.

KEY TO THE MALESIAN TAXA

The either cultivated that unnumberedtaxa are ones have not yet been observed to

African escape, or an one distinguished in this paperand includedhere for clarity's sake.

with of 1 a. Upper glume a marginal fringe relatively long, white hairs framing the

spikelet 2

b. Upper glume without such a frame 4

2a. Racemes 3—20, alternate. Spikelets paired 3

b. then the Racemes 2, subopposite, rarely 3, upper two subopposite. Spikelets

solitary 1. P. conjugatum R. de Koning & M. S. M. Sosef: Malesian species of Paspalum 289

3 a. Racemes 3-5(-9). Spikelets 1,8-2.5(-3.1) mm wide 2. P. dilatatum

urvillei b. Racemes (5—)10—20. Spikelets 1.2—1.5 mm wide 8. P.

4 a. Rachis 0.3—0.7 mm wide 5

b. Rachis more than 0.8 mm wide 6

hirsute all over P. 5 a. Spikelets 1.2—1.4 mm long. Upper glume . . 6. paniculatum

b. 2.5—3 the Spikelets mm long. Upper glume glabrous or pubescent along margins

only P. wettsteinii

6 a. Spikelets paired, at least in the middle of the raceme, one sometimes reduced

to minute glumes 7

b. Spikelets solitary 11

7 a. Inflorescence solitary from the uppermost sheath. Racemes 2—many. Lower

glume absent 8

b. Inflorescence 2—8 together from the uppermost sheath. Racemes 1 (or 2).

Lower glume developed P. decumbens

8a. Plants not robust, (10—)30—125(—150) cm tall. Blades 4—10 mm wide ... 9

Plants tall. Blades 10—34 wide P. b. robust, 100—200 cm mm virgatum

9 a. Spikelets broadly ovate to obovate, monochromate. Fertile lemma and palea

pale to brown, dull 10

b. Spikelets oblong to ovate or obovate, with a conspicuously dark centre and

paler margin. Fertile lemma and palea shiny dark brown P. plicatulum

10a. Racemes (3—)5—12(—16). Rachis (1.5—)2—3.5(—5) mm wide. Both spikelets

of the middle of the a pair usually well-developed in raceme. Upper glume

pubescent, very rarely glabrous 4. P. longifolium

b. Racemes 2—4(—6). Rachis 1—1.5(—1.9) mm wide. One of the spikelets of a

pair usually reduced. Upper glume glabrous 5. P. orbiculare

11a. Plantstoloniferous. Spikelets oblong, acute 12

b. Plants tufted, stolons absent. Spikelets ovate or obovate, rarely broadly so,

blunt 13

12a. Racemes 2, subopposite, rarely 3, then the upper 2 subopposite. Rachis spikel-

ed from the base. Lower glume present in at least some spikelets. Upper glume

pubescent 3. P. distichum

b. Racemes 2, strictly opposite, rarely up to 5, then the upper two strictly oppo-

site. Rachis at base partly without spikelets. Lower glume very rarely present.

Upper glume glabrous 9. P. vaginatum

13 a. Racemes 1—many, usually alternate. Spikelets 1.7—3.1 mm long. — Spikelets

and fertile lemma yellow brown to dark brown 14

b. Racemes 2, subopposite, rarely 3, then the upper 2 subopposite. Spikelets 3—4

— Fertile lemma P. notatum mm long. Spikelets green. yellow

14 a. Blades rarely hirsute all over. Peduncle glabrous. Spikelets broadly ovate to

broadly obovate 15

b. Blades usually hirsute all over. Peduncle usually sparsely hairy. Spikelets broad-

ly elliptic to ovate 7d. P. scrobiculatum var. horneri 290 BLUMEA - VOL. 30, No. 2, 1985

15 a. Spikelets usually suborbicular, brown to dark brown. Upper glume 5—13 -

nerved. Sterile lemma 5—9-nerved. Nerves concolorous. Fertile floret dark

brown in fruit 16

b. Spikelets usually broadly obovate, yellow-brown to brown. Upper glume and

sterile lemma 3- or 5-nerved. Nerves distinctly darker than the intervenium.

Fertile floret yellow-brown to brown in fruit 5. P. orbiculare

13 16a. Blades linear, 4—53 cm by 3—19 mm (index or more), narrowed towards

the base 17

b. Blades linear-lanceolate, 7-21 cm by 12-27 mm (index up to 10), distinctly

rounded at base. — Africa P. scrobiculatum var. lanceolatum

17 a. Spikelets 2.55—3.25 by 2.1—2.6 mm. Upper glume 7—13-nerved. Sterile lemma

7—9-nerved. — Racemes solitary or paired, then 1.5—7 cm long, or 3—8, then

8—15.5 cm long 18

b. Spikelets 1.8—2.6 by 1.45—2.15 mm. Upper glume and sterile lemma 5-7-

nerved. — Racemes (1 or) 2—6(—14), 1.5—9(—11) cm long

P. scrobiculatum 7c. var. bispicatum

18 Blades mm wide. Racemes 1 or 2, to 7 cm a. up to 9 up long

7a. P. scrobiculatum var. scrobiculatum

b. Blades 10—19 mm wide. Racemes 3—8, 8—15.5 cm long

7b. P. scrobiculatum var. auriculatum

1. Paspalum conjugatum Berg. — Fig. 1.

P. conjugatum Berg., Acta Helv. Phys.-Math. 7 (1762) 129, t. 8; Lamk., Tabl. Encycl. 1 (1791)

175; Kunth, Enum. I'l. 1 (1833) 51; Camus, Fl. Gen. I.-C. 7 (1922) 393; Ridley, Fl. Mai. Pen.

5 (1925) 218; Backer, Handb. Fl. Java 2 (1928) 120; Chase, Contr. U.S. Nat. Herb. 28, 1

(1929) 162, f. 105; Burk., Diet. Econ. Prod. 2 (1935) 1673;Reeder, J. Am. Arbor. 29 (1948)

297; Backer in Heyne, Nutt. PI. Indon. ed. 3, 1 (1950) 200; Schmid, l'Agron. Trop. 13 (1958)

304, f. 47-1; Metcalfe, Anat. Monoc. 1, Gram. (1960) 364; Bor, Grasses (1960) 336; Vickery,

Contr. N.S.W. Nat. Herb., Fl. Ser. 19,1 (1961) 117;Larsen, Dansk Bot. Ark. 20 (1963) 227;

Deshaprabhu, Wealt of India 7 (1966) 269; Hsu, Taiwania 13 (1967) 128; Monod de Froid. in

Backer & Bakh.f., Fl. Java 3 (1968) 570; Henty, Bot. Bull., Lae 1 (1969) 145; Gill., Rev. Fl.

Mai. 3 (1971) 180, pi. 24d, f. 37; Gould & Soderstrom, Canad. J. Bot. 52 (1974) 1084;

Beetle, J. Range Managem. 27 (1947) 347, f. 1 & 2; Powell in Paymans, New Guinea Veget.

(1976) 136, 167; Holm et al., World's Worst Weeds (1977) 134, f. 49 & 50; Hsu, Fl. Taiwan 5

(1978) 581, pi. 1439; J.W. Parham in A.C. Smith, Fl. Vit. Nov. 1 (1979) 336. - P. tenue

Gaertn., Fruct. Sem. PI. 2 (1791) 2, pi. 80, nom. superfl. - Lectotype: Rolander s.n. in

Herb. Berg. 36 (SBT, holo,n.v.; xerox in L, see note), Surinam.

For further Chase synonyms see (1929).

Perennials, not tufted, stoloniferous. Stolons leafy, rooting at the nodes,.glabrous, smooth. Culms ascending to erect, 40—80(—100) cm long, branching. Nodes of the stolons usually pubescent, the others glabrous. Sheaths slightly keeled, ciliate on the margins, pubescent on the outside collar; basal sheaths often pubescent, purplish, not reticulate, the higher ones otherwise glabrous, green. Ligule collar-shaped, 0.7—1.6 mm long. Blades flat, linear, narrowed at base, (2—)8—20(-23) cm by (2—)5—12(— R. de Koning & M. S. M. Sosef: Malesian species ofPaspalum 291

Fig. 1. Paspalum conjugatum Berg. a. Upper glume; b. sterile lemma (Darbyshire & Hoogland

7864).

3 white hairs the immediate base of the 15) mm, with some c. mm long at ligule,

less Peduncle sparsely pubescent on both sides, usually so beneath, to glabrous. 0.7—

1 mm diam., glabrous. Stalk up to 2 mm long, pubescent (hairs up to 4 mm long),

dark. Racemes 2, subopposite, rarely with a third below them, (5 —)8 —16 cm long.

0.6—1.2 Rachis with an adaxial, longitudinal groove, usually slightly curved down,

smooth Pedicel mm wide, spikeled from the base, margins to scabrous. terete, geni-

ovate culate, up to 4 mm long, smooth. Spikelets solitary, imbricate, (the lower ones often oblong), (1.5—)1.7 —1,8(—2.1) by (1.1 —)1.2— 1.4(—1.6) mm, pale green, apex acutish. Lower glume absent. Upper glume slightly convex, as long as the spikelet,

of 1 hairs the the bearing a fringe relatively long (c. mm) along margins, framing

spikelet, otherwise glabrous, nerves 2 (or 4), concolorous, midnerve suppressed.

not to Sterile lemma flat to slightly concave, glabrous, wrinkled, otherwise similar

as as the 0- or 3-nerved, the upper glume. Fertile lemma slightly convex, long spikelet, pergamentaceous, glabrous, smooth, pale green. Palea flat to slightly concave, 2-

otherwise lemma. Anthers 0.7 nerved, as the slender, c. mm long. Stigmas light brown to whitish (i. s.). Caryopsis broadly ovate, plano-convex, c. 1.1 mm long.

Distribution. A weed ofAmerican origin, introduced in the last halfof the 19th

century (not yet collected by e.g. Junghuhn), now common throughout Malesia.

Ecology. Not restricted to particular situations, though a preference for poor,

of shaded habitats has been reported. Often vegetation-forming. Present on a variety

soils, up to 1500 m alt. 292 BLUMEA - VOL. 30, No. 2, 1985

Vernacular names. English: buffalo grass (Malaya), carabao grass (Philippines),

sour paspalum; Sumatra: gujung tembagu, ujung abung, paitan burung, rumput kuda, r. paitan; Java: genjoran, jampang tjangah, jukut pait, kelemaran; Lesser Sunda Is-

lands: jampang pait; Borneo: blenna (Iban), rumput blanda; Philippines: bantotan

(Manobo), kauad-kauaran, kulape (Tagalog), lacatan (Cebuano), sacate (Bisayan);

also New Guinea: amdumaning (Eipo), sybsyby (Jali). For more names see Backer

of the (1950), who remarked that although some names indicate a bitter ('pait')

taste, this was to his own experience incorrect.

Collector's notes. Plants strongly branching and stooling, bright green, base

purple, stems sometimes purple pink, nodes white, bearded. Leaves dark green. Rachis

dark green. Inflorescence dull green. Spikelets yellow to yellow green. Anthers and

stigmas white or yellow. Fruits white to dark brown. Flowering at 11.00h. a.m. (de

Vogel 997).

Anatomy. See Metcalfe (1960).

Chromosome numbers. 2n = 20 (e.g. Hsu, 1967), 40 (e.g. Larsen, 1963), 80

(e.g. Gould & Soderstrom, 1974).

the of Phytochemistry. Deshaprabhu (1966) reported presence a haemostatic glucoside which reduces the time for the clotting of the blood by 50%.

Uses. A very important weed in crops, e.g. in sawahs, and plantations of bananas,

Malesia cocoa, coffee, papayas, pineapples, rubber, tea, etc. throughout (Holm et al.,

Different about the of this fodder Gener- 1977). opinions importance grass as a exist.

has but Backer stated that ally it a poor reputation, (1950) it is highly productive under good circumstances. Beetle(1974) reported that it is probably useful as fodder in areas of poor soil fertility, though suitable for grazing only when young since the fruits tend to stick in the throats of live-stock and choke them. Gianno (187, L) noted the same for ducks, geese, goats and other animals. Burkill (1935) noted that monkeys readily eat the grains. It is a valuable lawn grass due to its ability to form closed swards and its resistance to trampling.

to When wet, the fruit stick one's legs and other clothing and so may become very irritating (Gianno 187, L; Veldkamp, pers. comm.).

The Iban of Borneo use leaf concoctions for wound healing, while in the Sepik area (Papua New Guinea) crushed spikelets are used in the treatment of cuts, wounds and sores (Powel, 1976: 136). This is due to a haemostatic glucoside (see above).

The leaves are used by the Chimbu of Papua New Guinea for the lining of cooking ovens and for wrapping food(Powell, 1976: 167).

Notes. This species is reasonably variable especially in its vegetative characters probably due to ecological circumstances. No infraspecific taxa can be recognized.

The variety or forma ‘ristachya’ (Vanderyst, Bull. Agric. Congo Beige 9,1918, 245;

Beetle, 1974: 349, respectively) with three instead of two racemes in the inflores- cence occurs mixed with the normal form with two racemes in several collections, e.g. in the syntype ofP. conjugatum.

The type material of this name was not actually seen as the Bergius Herbarium

(SBT) does not send such specimens out on loan. Its Director, Dr. L.E. Kers, kindly sent us photocopies of the relevant material. There are two collections which can R. de Koning & M. S. M. Sosef: Malesian species of Paspalum 293

easily be identified with the present species, both annotated by Bergius. One was

collected by Swartz, presumably in the Caribbean, the other by Rolanderin Surinam.

The latter has a diagnosis written on it by Bergius and as only this provenance was

cited in the protologue, it seems the obvious lectotype.

Fig. 2. Paspalum dilatatum Poir. a. Upper glume; b. sterile lemma (Santos 5061).

2. Paspalum dilatatumPoir. — Fig. 2.

P. dilatatum Poir., Tabl. Encycl. 5 (1804) 35; Camus, Fl. Gen. I.-C. 7 (1922) 393; Backer, Handb.

Fl. Java 2 (1928) 120; Blatter & McCann, J. Bomb. Nat. Hist. Soc. 32 (1928) 640; Chase,

Contr. U.S. Nat. Herb. 28, 1 (1929) 169; Burk., Diet. Econ. Prod. 2 (1935) 1674;Backer in

Heyne, Nutt. Pi. Indon. ed. 3, 1 (1950) 201; Gardner, Fl. W. Austr. 1,1 (1952) 242, pi. 71;

Schmid, l'Agron. Trop. 13 (1958) 653; Bor, Grasses (1960) 338; Vickery, Contr. N.S.W. Nat.

Herb., Fl. Ser. 19, 1 (1961) 119; Deshaprabhu, Wealth of India 7 (1966) 269; Monod de

Froid. in Backer & Bakh.f., Fl. Java 3 (1968) 570; Henty, Bot. Bull., Lae 1 (1969) 146; B.C.

Stone, Micronesica 6 (1970) 216; Gill., Rev. Fl. Mai. 3 (1971) 183; Pohl & Davidse, Brittonia

23 (1971) 302; Gould & Soderstrom, Canad. J. Bot. 52 (1974) 1084;Kerguelen, Lejeunia n.s.

75 (1975) 222; Tsvelev, Grasses Soviet Union (1976) 668; Mehra & Chaudhary, Taxon 25

(1976) 633; Holm et al., World's Worst Weeds (1977) 358, f. 151 & 152; Hsu, Fl. Taiwan 5

(1978) 583, pi. 1440; J.W. Parham in A.C. Smith, Fl. Vit. Nov. 1 (1979) 3377. - Digitaria

dilatata Fl. Fr. 3 553. - Commerson Coste, (1906) Type: s.n. (Herb. Lamarck, P, n.v.;

photo in L), Argentine, Buenos Aires.

For further synonyms see Chase (1929).

Perennials, tufted, not stoloniferous. Culms ascending to erect, 40—120 cm long, rarely branching. Nodes glabrous or with some hairs. Sheaths slightly keeled; basal - 294 BLUMEA VOL. 30, No. 2, 1985

sheaths often pilose, purplish, not reticulate, the upper ones glabrous. Ligule bluntly triangular, up to 6 mm long. Blades flat, linear, narrowed at base, (5— )12—26(—39) cm by (3—)4-13 mm, usually with some up to 6 mm long hairs at the immediate base of the ligule, otherwise glabrous. Peduncle 0.8-1.3 mm diam., glabrous. Stalk

2.5(—7) mm long, appressed hirsute mixed with up to 9 mm long white hairs, brown.

Racemes 3—5(—9), alternate, the lowest 5—13 cm long, the others gradually shorter upwards. Internodes of main axis 0.3—0.7 times as long as the subtending raceme.

Rachis flat, winged, straight, frequently zigzag at the upper end, 1—1.3 mm wide, margins scabrous, spikeled to the base. Pedicels flattened, the inner ones 1.5—2.6 mm long, the outer ones shorter, edges ciliate. Spikelets paired, usually imbricate, ovate,

(2.8—)3—4 by 1.8—2.5(—3.1) mm, pale green to purplish, apex acute. Lower glume absent. Upper glume slightly convex, as long as the spikelet, faintly pilose and with a fringe of white, c. 2 mm long hairs along the margin framing the spikelet, nerves

5—7(—9), concolorous with to slightly darker than the intervenium, midnerve promi- nent. Sterile lemma flat, nerves 3—7, faintly pilose to glabrous, not wrinkled, other- wise as the upper glume. Fertile lemma slightly convex, weakly 3-nerved, thickly

Palea flat cartilaginous, glabrous, minutely papillose-striate, pale. to slightly con- cave, 2-nerved, otherwise as its lemma. Anthers slender, 1.2—1.5 mm long. Stigmas black about 2 to purplish (i. s.). Caryopsis suborbicular, plano-convex, up to mm long.

Distribution. Weed of American (sub)tropical origin, introduced in Malesia probably from Australia around 1900 (Singapore: 1902) as a cultigen, sometimes escaping: Java (Bogor, Jakarta, Priangan, Kedu, Malang, Besuki), Lesser Sunda

Islands (Flores, Timor), Borneo (Sabah), Philippines (Luzon), New Guinea (Central

Prov.).

Ecology. Roadsides, dry, damp or even water-logged sunny places, ash-grounds, wastegrounds, 50—2600 m alt.

Vernacular names. English: dallis grass (U.S., Philippines); Indonesian: rumput australi.

Collector's notes. Stigmas purplish black.

Chromosome numbers. 2n = 40, 60 (Gould & Soderstrom, 1974), 50 (Pohl &

Davidse, 1971), 54 (Mehra & Chaudhary, 1976).

Phytochemistry. See Deshaprabhu (1966) for an analysis.

Uses. troublesome weed banana and in the A in papaya plantations Philippines

(Holm et al., 1977).

fodder In some regions important as a highly productive grass though it requires careful management due to its minor competitive ability, especially on poor soil.

Suitable for grasslands at moderate height, therefore cultivated in Java above 1000 m, sometimes escaping, but setting seed badly and primarily propagated by cuttings

(Backer, 1950).

Due to its strong rooting habit it has been recommended as a suitable grass for the control of soil erosion (Deshaprabhu, 1966).

May cause diarrhoea in livestock possibly due to infections of an ergot, Claviceps paspali. R. de Koning & M. S. M. Sosef: Malesian species ofPaspalum 295

3. Paspalum distichum L. — Fig. 3.

P. distichum L., Syst. ed. 10, 2 (1759) 855; Benth., Fl. Austr. 7 (1878) 460; Ridley, FL Mai. Pen.

5 (1925) 218; Chase, Conti. U.S. Nat. Herb. 28, 1 (1929) 46; Gardner, Fl. W. Austr. 1 (1951)

241, 321; Metcalfe, Anat. Monoc. 1, Gram. (1960) 365; Bor, Grasses (1960) 338; Vickery,

Contr. N.S.W. Nat. Herb., Fl. Ser. 19,1 (1961) 119; Turpe, Lilloa 32 (1967)50; B.C. Stone,

Micronesica 6 (1970) 216; Jovet & Guedes, Taxon 21 (1972) 546; Fosberg, Taxon 26 (1977)

201; Guedes, Taxon 27 (1978) 128; Hsu, Fl. Taiwan 5 (1978) 583, f. 1441; J.W. Parham in

A.C. Smith, Fl. Vit. Nov. 1 (1979) 336; Renvoize & Clayton, Taxon 29 (1980) 339; Guedes,

Taxon 30 (1981) 301; Brummitt, Taxon 32 (1983) 281. - P. vaginatum Sw. var. fl Flugge,

Gram. Monogr., Paspalum (1810) 109. - Milium distichum Muhl., Descr. Gram. (1817) 78. -

Digitaria disticha Fiori & Paoletti, Icon. Fl. Ital. 111. 1 (1895) 16, f. 136. - Lectotype:

P. Browne in Herb. Linne 79-9, second fertile culm from the left (LINN, n.v.; microfiche

IDC), Jamaica. (See note on the confused typification.)

Digitariapaspalodes Michx., Fl. Bor. Am. 1 (1803) 46. — Milium paspalodes Elliott, Sketch Bot.

S. Carolina 1 (1816) 104. - P. michauxianum Kunth, Rev. Gram. 1 (1829) 25,nom. superfl.

- P. elliottii S. Warson in A. Gray, Man. Bot. ed. 6 (1890) 629, nom. superfl. - P. paspalodes

Scribner, Mem. Torrey Club 5 (1894) 29; Bor in Rechinger, Fl. Iran. 70 (1970) 494; Ker-

guelen, Lejeunia n.s. 75 (1975) 222; Vickery, Fl. N.S.W. Nat. Herb. 19, 2 (1975) 135;

Tsvelev, Grasses Soviet Union (1976)669; Srivastava, Acta Bot. Indica 10 (1982) 111, f. 1-7. - Anastrophus paspalodes Nash in Britton, Man. (1901) 75. P. distichum L. subsp. paspalo-

des Thell., Mem. Soc. Nat. Cherbourg 38 (1912) 77. - Type: Herb. Michaux (P, n.v.),

U.S.A., South Carolina.

P. distichum L. var. longirepens Domin, Bibl. Bot. 85 (1915) 289. - Type: Wools ex Herb.

F. von Mueller, anno 1877 (K), Australia, Sydney, Manley Beach.

P. distichum L. var. microstachyum Domin, Bibl. Bot. 85 (1915) 289. - Type: Domin s.n.,

anno 1910 (PR, holo,n.v.), Australia, Queensland.

For further synonyms see Chase (1929).

Perennials stoloniferous, sometimes tuft-

ed. Stolons leafy, rooting at the nodes,

glabrous, smooth. Culms ascending to

erect, 9-60(-100) cm long, rarely

branching. Nodes often with some c. 3

mm long, white hairs. Sheaths slightly

keeled, with some c. 2.5 mm long, white

hairs along the margins, at least at the

apex, usually in the upper third, other-

wise glabrous, the basal ones not reticu-

late. Ligule collar-shaped, 0.4—0.8 mm

long. Blades flat, linear, narrowed at base,

(1 —)6—10( —20) cm by (1.5—)3—5(—9)

mm, usually with some c. 2.5 mm long,

white hairs at the immediate base of the

ligule, otherwise glabrous, rarely pubes- Fig. 3. Paspalum distichum L. a. Upper glume; cent. Peduncle 0.7—0.8 mm diam., gla- b. sterile lemma (PNH 33167).

brous or with a few, white hairs. Stalk up

to 1 mm long, green-brown, with some c. 2.5 mm long, white hairs. Racemes 2, sub-

with below opposite, rarely a third one them, (2-)3—5(-8) cm long. Rachis flatten- 296 BLUMEA - VOL. 30, No. 2, 1985

ed, straight, not winged, 1.2-2 mm wide, spikeled from the base, margins scabrous.

the Pedicels flattened, straight, 0.2—0.5 mm long, ciliate. Spikelets solitary, upper

ones usually imbricate, oblong, 2.9—3.4(— 4) by 1.3—1.6(—1.8) mm, pale green, apex acute. Lower glume at least in some spikelets present, reduced to a minute

flat, 3- scale, rarely more developed, then oblong, up to 2 by 1 mm, acute, 1- or

nerved. Upper glume slightly convex, as long as the spikelet, pubescent, sometimes

concolorous obscurely so, nerves (3 or) 5 or 7, with the intervenium, midnerveprom-

the inent. Sterile lemma flat, glabrous, not wrinkled, as upper glume. Fertile lemma

convex, 0- or weakly 3-nerved, pergamentaceous, smooth, apex with some short,

Anthers stiff cilia, pale green. Palea flat, 0- or weakly 2-nerved, as the lemma. slender,

c. 1.5 mm long. Stigmas dark brown to purple (i.s.). Caryopsis ovate, plano-convex,

c. 1.5 mm long.

Distribution. A pantropical weed, probably introduced in Malesia: Malaya

(Penang), Java (Priangan), Philippines (Luzon), Moluccas (Buru).

Ecology. Wet marshlands, swamps, in polluted, shallow water, along irrigation

ditches, etc., up to 1725 m alt.

Vernacular names. English: ginger grass, knot grass,victoria grass, water couch;

Indonesia: rumput italia; Philippines: gagayut (Wfumo).

Collector's notes. Glaucous, greenish withblack anthers and styles.

Anatomy. See Metcalfe (1960).

Chromosome numbers. 2n = 40,48, 60 (e.g. Turpe, 1967).

Cytology. Srivastava (1982, as P. paspalodes) reported obligate apomixis for

Indian representatives.

Uses. Considered to be a valuable pasture grass on alluvial flats and useful as a

soilbinder on stream banks. Sometimesfound as a weed in sawahs (Bor, 1960).

Notes. When Linne described P. distichumhe mentionedmaterial sent to him by

P. Browne. Bor (Fl. Iraq 9, 1968, 494) without any explanation indicated that the

name had been misapplied and that the correct name would be P. paspalodes (Michx.)

Scribner (‘spaloides’). In 1970 he explained that the specimen on sheet 79.9 in the

Linnean herbarium (LINN) which most closely agreed with the Linnean description

was a grass which until then had been known as P. vaginatum Sw. This name ought

therefore to be reduced into the synonymy of P. distichum, while the species com-

monly called by that name needed a new one.

The sheet 79.9 in the Linnean herbarium bears one sterile and three fertile culms.

Linne had marked them with 'Br.' indicating that they represent P. Browne collec-

the sheet Guedes tions. Unfortunately contains a mixture as was observed by Jovet &

the the P. the (1972): culm on right-hand side is indeed vaginatum, but two others, however, are something quite different.

From Linne's original description ('spicibus duabus, altera subsessili', i.e., only one of the two spikes subsessile) Jovet & Guedes argued that thatreferred to the two left- hand culms since the right-hand one has both racemes equally subsessile. The latter author therefore retypified P. distichum choosing the fertile culm second from the left whereby both names were restored to their classical usage. Fosberg (1977) dis- agreed with this, remarking that 'Br.' had been written closest to the right-hand culm R. de Koning & M. S. M. Sosef: Malesian species ofPaspalum 297

which in his opinion indicated that Linne had thus wantedto indicatethat this was the

'true' specimen of P. Browne, whereby Bor's original lectotypification ought to be fol-

lowed. Guedes (1978), in his turn, countered this view and argued that the relative po-

of should be much while would be sition a mark not given so weight, it impossible to

that all not of the Brownean col- prove anymore specimens were already part same that lection originally received by Linne. In our opinion he correctly pointed out

types should be selected conform the original descriptions and preferably in accor-

choose of dance with established use ('so we a specimen everybody's P. distichum’).

To put an end to all this confusion Renvoize & Clayton (1980) then proposed

that the combinationP. distichum should be rejected, whereby P. vaginatum would be restored and P. distichum would become P. paspalodes. Guedes (1981) retorted

'such classical should dismissed.' to this new viewpoint that a very name not be

nomenclature end all A vote of the Committee on (Brummitt, 1983) put an to

these squabbles and resulted in a vindication of Guedes' typification. So we have P.

distichum the second from the left Herb. Linne L. being typified by specimen on

sheet P. Sw. the Swartz herbarium. 79-9 and vaginatum typified by a specimen in

4. Paspalum longifolium Roxb. — Fig. 4.

P. longifolium Roxb. [Hort. Beng. (1814) 7, nomen] Fl. Ind. 1 (1820) 283;Trin., Diss. Bot. Alt.

(1826) 109; Sp. Gram. Icon. 2 (1829) t. 138; Buse in Miq., Pi. Jungh. 3 (1854) 383; Miq., Fl.

Ind. Bat. 3 (1857) 432; Camus, Fl. Gen. I.-C. 7 (1922) 392; Ridley, Fl. Mai. Pen. 5 (1925)

217; Boi, Fl. Assam 5 (1940) 253; Jansen, Reinwardtia 2 (1953) 321; Schmid, l'Agron. Trop.

13 (1958) 302, f. 47-2; Bor, Grasses (1960) 339; Henty, Bot. Bull., Lae 1 (1969)246; Gill.,

Rev. Fl. Mai. 3 (1971) 182, pi. 24a, f. 14; Hsu, Taiwania 16 (1971) 292; Malik & Tripathy,

Broteria 41 (1972) 55; Koyama in Walker, Fl. Okinawa (1976) 216. - Type : Roxburgh s.n.

(BM, holo, photo in K), India, cult.

P. scrobiculatum L. var. 0 Flugge, Gram. Monogr., Paspalum (1810) 88. - Type: Herb. Klein

(ubi?), isotype of P. flexuosum Presl?

P. sumatrense Roth ex R. & S., Syst. Veg. 2 (1817) 316; Roth, Nov. PI. Spec. (1821) 35. -

Type: Heyne s.n. (B, holo,lost; K, p.p.), East Indies.

P. flexuosum Klein ex Presl, Rel. Haenk. 1 (1820) 216; Miq., Fl. Ind. Bat. 3 (1855) 433. - Type :

microfiche first sheet from Klein in Herb. Willdenow no. 1574,p.p. (B, holo, n.v.; IDC!: left;

isotype of P. scrobiculatum var. (3 Flugge?), India.

— P. Steudel Fl. Ind. P. cognatum Steudel, Syn. 1 (1853) 28. zollingeri var. (3gracilior Miq., Bat.

3 (1857) 431. - Type : Zollinger 192 (P, holo, n.v.; B, BM, K, L), Java.

De P. houttuynii Van Hall ex Vriese, PI. Ind. Bat. Or. 2(1857)113.-Type: Kleinhoffin Herb.

Reinwardt (?L, holo, but not found), Java.

P. longifolium Roxb. var. hirsutum Boerl., Ann. Jard. Bot. Btzg. 8 (1890) 49. - Ty p e : Boerlage

s.n. (BO, holo), Java, Bogor, Botanic Garden, 6 October 1896.

P. scrobiculatum L. vai. philippinense Merr., Philip. J. Sc. 1 (1906) Suppl. 345. - Lectotype :

BS 1449 (Ramos) (PNH, holo; BO), Philippines, Luzon, Rizal Prov., Morong, August 1906.

3 (1908) 167. - P. longifolium Roxb. var. trichocoleum Hackel, Philip. J. Sc. Bot. Type:

Merrill 5455 (W, holo, PNH, lost, n.v.; BO, L), Philippines, Mindanao, Surigao Prov., Caraga,

6 October 1906.

P. platycoleum Ridley, Fl. Mai. Pen. 5 (1925) 217. - Type : SF4509 (Nur) (SING, holo; K),

Selangor, Port Swettenham, 13 February 1919.

P. orbiculare Forst.f. var. otobedii Fosb. & Sachet, Micronesica 18 (1984) 83. - Type : Otobed

PW-10090 (US, holo), Carolines, Palau I., Koror I., 22 August 1977. 298 BLUMEA - VOL. 30, No. 2, 1985

Fig. 4. Paspalum longifolium Roxb. a. Upper glume;b. sterile lemma (Ramos 1956).

Perennials, tufted, not stoloniferous. Culms erect to ascending, (20—)50 — 125

(—150) cm long, rarely branching. Nodes glabrous or stiffly hairy. Sheaths slightly keeled, the basal ones hirsute at base or glabrous, purplish, rarely somewhat reticu- late, the higher ones smooth or with a pilose margin, otherwise glabrous. Ligule col- lar-shaped, 1—3 mm long. Blades usually once folded lengthwise, linear, narrowed towards the base, (2—)20—40(—56) cm by (2—)4—9(—9.5) mm, appressed hirsute at least behind the ligule or at base. Peduncle (0.6—)0.7—1.5 mm diam., glabrous or with a few hairs. Stalk 0.8—3 mm long, appressed hirsute, mixed with some white, up to 11 mm long hairs to scabrous, black. Racemes (3 —)5—12( — 16), alternate, 1.5—8

cm long. Internodes of the main axis 0.15—0.8 times as long as the subtending ra- ceme. Rachis flat, straight, winged, (1.5—)2—3.5(—5) mm wide, spikeled to the base, margins scabrous. Pedicels terete to slightly quadrangular or 2-edged, 0.5—1.5 mm long, scabrous all over or only along the edges, the inner one usually shorter than the outer one. Spikelets paired (the inner spikelet sometimes reduced to minute glumes, but usually well-developed at least in the middle part of the raceme), imbricate, broadly ovate to broadly obovate, 2.1—2.5(—2.8) by 1.2—2(—2.2) mm, yellow-green to light brown, sometimes purplish, apex acuminate. Lower glume absent. Upper

the all the glume convex, as long as spikelet, sparsely pubescent over or only along margins, very rarely entirely glabrous, nerves 3, darker than the intervenium, mid- nerve prominent. Sterile lemma flat to slightly convex, not wrinkled, as the upper glume. Fertile lemma convex, 0—5-nerved, thickly cartilaginous, usually glabrous, very rarely sparsely pubescent, or the margins with a few hairs, papillose-striate, pale brown. Palea slightly convex, 0- or 2-nerved, as the lemma. Anthers slender, 0.7—1.1 mm long. Stigmas brown to purplish black (i.s.). Caryopsis broadly obovate, unequal- ly biconvex, c. 1.5 mm long.

Distribution. India (Assam), Nepal, Sri Lanka to Vietnam, to the Pacific (Ryu- kyu I., Samoa) and N. Australia (Queensland), common throughout Malesia. R. de Koning & M. S. M. Sosef: Malesian species of Paspalum 299

Ecology. Solitary or in groups in moist places, e.g. along riverbanks, in swamps,

in to 60 water. bogs and pools, floating grass communities, growing in up cm deep A

common invader in wet and open, disturbed places. Weed in sawahs, associated with e.g. Bothriochloaglabra, Echinochloa, Eriocaulon, Hymenachne, Miscanthus, Oryza.

alluvial On sandy, loamy, clayey, soils, up to 1700 m alt.

2 Vernacular names. Malaya: paha belalang, rumput patah siku; Riouw: siak ;

Sumatra: gereman badah(Ulunese); Java: jukut cariang, rumput jilurang, wawaderan;

Borneo: cakan belangkas; Celebes: bulibuliba merah; New Guinea: wabiwabita

(Minufia, Kabubu), waiofana (Onjob, Naukwata).

Collector's notes. Erect to ascending, occasionally rooting from the lower

rhizomes whole leaves red- nodes, no or stolons, plant green or occasionally purplish,

Inflorescence Rachis dish green to green. green. purple. Racemes green, greenish brown, reddish. Spikelets purplish when flowering, dark brown. Anthers yellow,

black. black. Fruit brown. orange, Stigmas purple, greenish,

Chromosome numbers. 2n=40 (e.g. Hsu, 1971), 50 (e.g. Malik & Tripathy,

1972).

Uses. Some value fodder for buffaloes as a grass water (Bor, 1960).

Notes. Paspalum scrobiculatum L. var. philippinense Merr. has here been lecto-

typified with BS 1449 (Ramos), as this was the only collectionof the three cited by

Merrill which we have seen.

Plants with a hirsute sheath have been described as var. hirsutum by Boerlage

(1890), but the pubescence is too variable to base distinct taxa upon.

form with the rachis Bor (1940, 1960) mentioned a var. lorirachis Bor, a robust

8 such have been and the status must in doubt. up to mm wide. No plants seen remain

5. Paspalum orbiculare Forst. f. — Fig. 5.

P. orbiculare Forst.f., Fl. Ins. Austr. Prodr. (1786) 7;Camus, Fl. Gen. I.-C. 7 (1922) 391; Ridley,

Fl. Mai. Pen. 5 (1925) 217; Burk., Diet. Econ. Prod. 2 (1935) 1674;Schmid, l'Agron. Trop.

13 (1958) 302; Bor, Grasses (1960) 340; Vickery, Contr. N.S.W. Nat. Herb., Fl. Ser. 19, 1

(1960) 118; Henty, Bot. Bull., Lae 1 (1969) 146; Nath et al., Cytologia 35 (1970) 111; Gill.,

Rev. Fl. Mai. 3 (1971) 184, pi. 24b, f. 15; Hsu, Taiwania 16 (1971) 292; Khosla & Mehra,

Taxon 22 (1973) 560; J.W. Parham in A.C. Smith, Fl. Vit. Nov. 1 (1979) 338; Dujardin,

Canad. J. Bot. 57 867. - P. scrobiculatum var. Gram. (1979) y Flugge, Monogr., Paspalum

Bot. - (1810) 88. - P. scrobiculatum L. var. orbiculare Hackel, Jahrb. 6 (1885) 233. Type :

Forster s.n. (GOET, holo, n.v.; K), Society Islands.

P. longifolium Roxb. var pseudo-orbiculare Jansen, Reinwardtia 2 (1953) 321. - Type : Santos

5 (MICH, holo), Philippines, Luzon, Mountain Prov., Baguio, 28 March 1940.

Perennials, tufted, not stoloniferous. Culms erect, (10—)30—70(—100) cm long, rarely branching. Nodes glabrous, rarely with a fewhairs. Sheaths not keeled, glabrous,

the basal ones sometimes hirsute in the lower half, not reticulate. Ligule collar-

shaped, 0.3-l(-2) mm long. Blades flat or once folded lengthwise, linear, narrowed

towards the base, (2-)8-25(-40) cm by (1.5—)4—7(—8.5) mm, hirsute at base at

least behind the ligule. Peduncle 0.5-0.7(-0.8) mm diam., glabrous. Internodes of

the main axis 0.25—1 times as long as the subtending raceme. Stalk 0.5—1.5 mm - 300 BLUMEA VOL. 30, No. 2, 1985

b. Fig. 5. Paspalum orbiculare Forst. f. a. Upper glume; sterile lemma; c. fertile lemma and its palea (Koorders 42216).

long, appressed hirsute usually mixed with one or more up to 7 mm long, white hairs, black. Racemes 2—4(—6), alternate, 1—6.5 cm long. Rachis flat, winged, straight to

slightly zigzag, 1—1.5(—1.9) mm wide, spikeled from the base, margins scabrous.

Pedicels oval diam. 0.3—0.8 scabrous. in or 2-edged, mm long, edges Spikelets paired or solitary, the inner one sometimes developed, usually reduced in the middle of the raceme, sometimes absent at its ends, imbricate, usually broadly obovate, less often suborbicular to broadly obovate, (1.7 —)1.9—2.3(—2.7) by (1.2—)1.5—1.8(—2) mm, yellow-brown to brown, apex acute to rounded. Lower glume absent. Upper glume convex, as long as the spikelet, glabrous, nerves 3 or 5, darker than the intervenium, midrib prominent. Sterile lemma slightly convex, not wrinkled, as the upper glume.

Fertile lemma convex, 0-, or 3-, or 5-nerved, coriaceous, glabrous, papillose-striate, yellow-brown to brown. Palea slightly convex, 0- or 2-nerved, as its lemma. Anthers slender, 0.5-0.9 mm long. Stigmas dark purple to black (i.s.). Caryopsis broadly ovate to broadly obovate, unequally biconvex, c. 1.3 mm long.

Distribution. India (Assam), Sri Lanka, Nepal, to Taiwan, Hongkong to Poly- nesia (to Hawaii, New Caledonia) and N. Australia (Queensland, New South Wales), throughout Malesia.

Ecology. Generally in moist places, e.g. marshes, swamp grasslands, ponds, also in sunny grassy places near the sea, on sandy soils, clay, or peat, in up to 25 cm deep water. Associated with Cyperus, Eleocharis, Ophiuros, Themeda, Xyris. Up to 2200 m altitude.

Vernacular names. None recorded on the sheets, often confused with P. scro- biculatum s.l., q.v.

Collector's notes. Culms coarse, erect, ascending with an angle, tufted, thick

soft at and base, grey-green, branching horizontally. Leaves milky green, dullgreen.

Spike pinkish, green, brown. Spikelets pale green, green-brown. Anthers black. Stig- mas purple. Fruit green. R. de Koning & M. S. M. Sosef: Malesian species ofPaspalum 301

numbers. 2n 20 40 Khosla & Chromosome = (Dujardin, 1979), (e.g. Mehra,

1973), 54 (Nath et al„ 1970), 60 (Hsu, 1971).

Uses. None recorded, but cf. P. scrobiculatum.

in Notes. This species may cause difficulties the identification when the inner

spikelets are all reduced whereby the outer ones may appear to be solitary. Especial-

ly P. scrobiculatum var. bispicatum is then very similar. The spikelets in P. orbiculare

discolorous and are usually broadly obovate with 3 or 5 nerves in the upper glumes

sterile lemmas, while in var. bispicatum the spikelets are usually suborbicular and the

concolorous. nerves are 5 or 7,

The forma depauperata Chase mistakenly included by her in P. longifolium repre-

form of the and needs taxonomical distinc- sents a depauperate present species no

tion.

A peculiar collection is Hosokawa 5959 (L) from Leval, Ponape. It has tumid

spikelets with the fertile lemmas nearly completely enveloping the palea, a situation

not encountered elsewhere.

6. Paspalum paniculatum L. — Fig. 6.

P. paniculatum L., Syst. Nat. ed. 10, 2 (1759) 855; Stapf in Prain, Fl. Trop. Afr. 9 (1920) 577;

Chase, Contr. U.S. Nat. Herb. 28, 1 (1929) 122, f. 72; Metcalfe, Anat. Monoc. 1, Gram.

(1960) 366; Vickery, Contr. N.S.W. Nat. Herb., Fl. Ser. 19,1 (1961) 121; Henty, Bot. Bull.,

Lae 1 (1969) 146; Bosser, Mem. Orstom 35 (1969) 399, f. 149e & f; Davidse & Pohl, Canad.

J. Bot. 50 (1972) 275; J.W. Parham in A.C. Smith, Fl. Vit. Nov. 1 (1979) 335. - Panicum

paniculatum O. Kuntze, Rev. Gen. PI. 3 (1898) 363. - Type: P. Browne in Herb. Linne

79-11 (LINN, holo, n.v.; microfiche IDC!), Jamaica.

For further synonyms see Chase (1929).

Perennials, tufted, not stoloniferous. Culms ascending to erect, 35—100(—120)

hairs. cm long, often branching. Nodes pilose with up to 5 mm long, white or brown

Sheaths slightly keeled, hirsute, rarely glabrous, the basal ones not reticulate. Ligule

collar-shaped, c. 3 mm long. Blades flat, linear, rounded at base, (4—)20—35 cm by

(5—) 10—25 mm, hirsute, rarely glabrous, with a tuft of long, white hairs at the im-

mediate base of the ligule. Peduncle 0.9—1.3 mm diam., with scattered hairs. Stalk

up to 2 mm long, hirsute, mixed with

some up to c. 5 mm long, white, some-

times flattenedhairs, dark brown..Racemes

10—40(-60), alternate, the lowermost

5.5-8 cm long, the others distally becom-

ing shorter. Internodes of the main axis

0.1—0.3 the times as long as subtending.

Rachis flat, winged, straight to slightly zig-

zag at the upper end, 0.3-0.5 mm wide,

spikeled from the base, margins scabrous

and often with some hairs. Pedicels terete Fig. 6. Paspalumpaniculatum L. a. Upper glume; b. sterile lemma (Brass 28208). to 2-edged, the outer ones 0.6—1.2 mm 302 BLUMEA - VOL. 30, No. 2, 1985

scabrous. sometimes few soli- long, the inner ones shorter, edges Spikelets paired, a

within at least the ones imbricate, obo- tary or rarely ternate a single raceme, upper

vate, 1.2— 1.4 by 0.9—1 mm, yellow-brown or purplish with brown dots, apex round- ed. Lower glume absent. Upper glume convex, slightly shorter than, sometimes as

long as the spikelet, shortly appressed hirsute, nerves 5, concolorous with to slightly

darker than the intervenium, midnerve prominent. Sterile lemma flat, hirsute along

the margins, 3- or 5-nerved, not wrinkled, as the upper glume. Fertilelemma convex,

0- or 3nerved, thickly coriaceous, glabrous, smooth, yellow. Palea flat to slightly con-

vex, 2-nerved, as the lemma. Anthers slender, c. 1 mm long. Stigmas dark brown to

dark purple (i.s.). Caryopsis ovoid, unequally biconvex, c. 1.2 mm long.

Distribution. Tropical South America and West Africa, introduced elsewhere in

the tropics, e.g. in Malesia: New Guinea (Vogelkop, Jayapura, West Sepik,East Sepik,

Eastern Highlands, Southern Highlands, Central, Northern, Morobe, Milne Bay, New

Britain).

Ecology. Locally common along roadsides, wastegrounds, in regrowth ofAlbiz-

zia, coconut plantations, rocky stream banks, etc. on loamy clay or volcanic sands,

up to 1400 m alt.

Vernacular names. English: galmarra grass, Russell river grass; New Guinea:

hanu (Kutubu), samara (Orokaiva, Mumuni), wolo (Miwaute, Wapi).

Collector's notes. Culms erect. Sheaths slightly viscid, the lower with irritant

below. hairs, hairs very sharp. Leaves dull green above, lighter Spikelets purplish.

to adhere to animals and thus Fruit brownish, grey, tending man during rain and transported (Bosser, 1969).

Anatomy. See Metcalfe (1960).

Chromosome numbers. 2n = 20,40 (e.g. Davidse & Pohl, 1972).

Uses. A weed of cultivated land. Introduced as a pasture grass, but apparently

not palatable to cattle (Parham, 1979).

Notes. Schodde 2153 (K, L) from Papua, S. Highlands, Lake Kutubu, has ternate

spikelets, the third ones much reduced.

7. Paspalum scrobiculatum L. — Diagram 1 a—c (see p. 285).

For the references see under the varieties.

a. var. scrobiculatum

P. scrobiculatum L., Mant. 1 (1767); R. & S., Syst. Veg. 2 (1817) 382; Buse in Miq., PI. Jungh.

(1854) 382; Miq., Fl. Ind. Bat. 3 (1857) 430; Camus, Fl. Gen. I.-C. 7 (1922) 391, t. 39, f. 5 &

6; Burk., Diet. Econ. Prod. 2 (1935) 1675; Henr., Blumea 3 (1940) 440; Bor, Grasses (1960)

340; Deshaprabhu, Wealth of India 7 (1966) 270; Henty, Bot. Bull., Lae 1 (1969) 145; Gill.,

Rev. Fl. Mai. 3 (1971) 185, f. 38; Hsu, Taiwania 16 (1971) 292; Clayton, Kew Bull. 30

(1975) 101. - P. scrobiculatum L. vai. frumentaceum Stapf in Prain, Fl. Trop. Afr. 9 (1920)

575, nom. superfl. - Lectotype: Herb. Linne no. 79-4 (LINN, holo, n.v.; microfiche

IDC!), India orientalis.

[P. frumentaceum Rottl.ex Beauv., Agrost. (1812) 10,171, nom.inval.,'Indiaorientalis','Rottb.'] R. de Koning & M. S. M. Sosef: Malesian species ofPaspalum 303

P. scrobiculatum L. var. turgidum Buse in De Vriese, PI. Ind. Bat. Or. 2 (1857) 113. - P. com-

mersonii Lamk. var. turgidum Jansen, Reinwardtia 2 (1953) 320. - Type: Reinwardt 66

(L, holo), Java.

Plants perennial (annual?), tufted, not stoloniferous.Culms ascending to erect, 8—

20(—55) cm long, sometimes branching. Nodes glabrous, rarely setose. Sheaths not

keeled, the basal ones usually glabrous, sometimes hirsute all over, not reticulate, the

cauline ones usually hairy along the margins or only in the upper part, rarely gla-

brous. Ligule collar-shaped, 0.3—1.8 mm long. Blades flat or once foldedlengthwise,

linear, narrowed towards the base, (4-)7—20(—30) cm by (3—)5—7(—9) mm, hir-

sute at base along the margins and above or at least behind the ligule, rarely hirsute

all over. Peduncle 0.5-1 mm diam., glabrous. Stalk 0.8-2 mm long, appressed hir-

sute, sometimes mixed with white, up to 5 mm long hairs, black. Racemes solitary,

terminal, or paired, subopposite or alternate, 1.5—7 cm long. Internodes of the main

axis up to 0.25 times as long as the subtending raceme. Rachis flat, straight to slight-

from the ly zigzag, winged, (0.8—)1.3—2.2(-3) mm wide, spikeled base, margins

Pedicels 0.4—1.3 scabrous. straight, terete to 2-edged, mm long, edges scabrous.

less often Spikelets solitary, imbricate or not, usually suborbicular, broadly ovate to

obovate, 2.55—3.25 2.1—2.6 brown dark brown, rounded broad- by mm, to apex to

ly, shortly, bluntly acuminate. Lower glume absent, rarely more or less developed.

Upper glume strongly convex, as long as the spikelet, glabrous, nerves 7,9, 11, or 13,

concolorous with the intervenium, midnerve prominent. Sterile lemma flat to slightly

the convex, 7- or 9-nerved, usually wrinkled along margins, occasionally cartilaginous

Fertile lemma or partly so. strongly convex, 0-, 3-, or 5-nerved, coriaceous, glabrous,

at Palea the papillose-striate, dark brown maturity. slightly convex, 0- of 2-nerved, as

lemma. Anthers slender, 0.8—1.2 mm long. Stigmas white or purple (i.s.). Caryopsis

broadly ovate to obovate, usually biconvex, c. 2 mm long.

Distribution. World distribution uncertain due to doubtful identifications in

the literature, seen from India (throughout), Sri Lanka to the Pacific (New Hebrides),

Malesia: Java (Bantam, Jakarta, Ceribon, Pekalongan, Banyumas, Kedu, Semarang,

Japara-Rembang, Madiun, Kediri, Surabaya, Malang, Madura, Kangean I.), Lesser

Sunda Islands (Sumba, Flores, Timor), New Guinea (see note). Not seen from Africa.

Ecology. Roadsides, sawahs, teak plantations, usually in moist places on clayey

soils, loam or volcanic sands, up to 650 m altitude. Exact data are difficult to verify because of the confusion with the other varieties.

Vernacular names. English: kodo millet, kodra millet; Java: rebu bawang, rum-

2 put kinangan; Madura: rebbha ghung mloko; suket krisik (Tayu).

Chromosome numbers. None recorded, but see the introduction and var. bi-

spicatum.

Uses. Cultivated for the grains, but reported to be poisonous possibly due to in-

fections of the outer husks by Sorosporium paspali and Uredo paspali-scrobiculari.

'Eaten with caution, except when the eaters deliberately set themselves to get intoxi-

cated' (Burkill, 1935). Bor (1960) mentioned that elephants which had raided the

crop died as a result of their depredations. Storage of more than six months seems to remove the poisonous principle (Deshaprabhu, 1966). 304 BLUMEA - VOL. 30, No. 2, 1985

Notes. This has been described annual. The have variety as an specimens we seen,

however, when they had a root system, were invariably perennial. As the roots were

often absent we cannot exclude the possibility of an annual cycle, but it seems un-

likely.

Beauvois (1812) mentioneda Paspalum frumentaceum 'Rottb.' (no doubt a mis-

take for because the from found 'Rottler' of provenance India), a name probably on

a label. Although the name appears in various reports, e.g. Roemer & Schultes (1817)

('Rottler'), the combination was never validly published as it was generally cited as a

synonym of P. scrobiculatum. Stapf (1920) used the epithet at the varietal level

under that name without citing a source. As he stated that this form represents the

still typical variety his combination is invalid, as the autonym must now be used. His

specimens (inch Rottler s.n.., K) indeed belong to var. scrobiculatum.

The variety turgidum Buse (1854) was proposed for small perennials with few

racemes and large spikelets. Again the autonym is required.

According to Henty (1969) several introductions have been tried in Papua New

Guinea, but none seem to have persisted.

b. var. auriculatum (Presl) Merr.

P. scrobiculatum L. var. auriculatum (Presl) Merr., Philip. J. Sc. 1 (1906) Suppl. 345. - P. auricu-

latum Presl, Rel. Haenk. 1 (1830) 217; Miq., Fl. Ind. Bat. 3 (1857) 432; Henr., Blumea 3

(1940) 440. - Lectotype: Haenke s.n. (PR, holo; K), Philippines, Luzon, Sorzogon (see

note).

P. zollingeri Steudel, Syn. 1 (1853) 28; Miq., Fl. Ind. Bat. 3 (1857) 431; Camus, Fl. Gen. I.-C. 7

(1922) 391; Schmid, l'Agron. Trop. 13 (1958) 302. - Type: Zollinger 48 (P, holo, n.v.; B, K, L), Java.

Differs from the typical variety by:

Culms to 135 Cauline sheaths Blades up cm long. glabrous. 20-40(—53) cm by

10—16(—19) mm. Peduncle 1—1.9 mm diam. Racemes 3—8, alternate, 8—15.5 cm

7-nerved. Sterile lemma long. Upper glume 7- or 9-nerved, usually more or less

smooth.

Distribution. General distribution uncertain because of the confusions in iden-

tity. Cultivated and escaping in at least India, Sri Lanka, Thailand and Malesia: Java

(Bantam, Kedu), Lesser Sunda Islands (Flores, Alor, Timor), Philippines (Palawan,

Paragua, Leyte), Moluccas (Sula).

Ecology. Wet monsoon forests, grassplains, along roads on damp open soil on

loam, soil, to 650 altitude. yellow mecamie up m

Collector's notes. None recorded.

Chromosome numbers. Not known with certainty.

Uses. A cereal in India, sown in June, harvested in November(Panigrahi 20846,L)

Notes. The type collection in PR is a mixture of three culms mounted on the righthand side belonging to the present variety and one culm on the left side belong-

to sterile ing var. bispicatum (with indurated lemmas). Presl's description agrees with

the first three which therefore be must regarded as type material. R. de Koning & M. S. M. Sosef: Malesian species ofPaspalum 305

Fig. 7A. Paspalum scrobiculatum L. var. bispicatum Hackel. a. Upper glume; b. sterile lemma

(Hoogerwerf227).

Hackel — c. var. bispicatum Fig. 7A.

L. P. scrobiculatum var. bispicatum Hackel in Kneucker, Allg. Bot. Zeitschr. 20 (1914) 146. -

Type: Kneucker Exsicc. 803 (Merrill & McGregor) (W, holo, n.v.; BM, K, L, UPS), Philip-

pines, Luzon, Rizal Prov., San Petro Macati, 10 m alt., July 1910.

P. commersonii Lamk., Tabl. Encycl. 1 (1791) 175, t. 43; Ridley, Fl. Mai. Pen. 5 (1925) 218;

Jansen, Reinwardtia 2 (1953) 319; Metcalfe, Anat. Monoc. 1, Gram. (1960) 362; Bor, Grasses

(1960) 335; Tateoka, Amer. J. Bot. 52 (1965) 167; Henty, Bot. Bull., Lae 1 (1969) 145; B.C.

Stone, Micronesica 6 (1970) 218. - P. scrobiculatum L. var. commersonii Stapf in Prain, Fl.

Trop. Afr. 9 (1920) 573; Camus, Fl. Gen. I.-C. 7 (1922) 391. - Type: Commerson s.n.

(P, holo; L), Mauritius.

P. kora Willd., Sp. PI. ed. 4,1 (1797) 332; Roxb., Fl. Ind. ed. Carey 1 (1832) 279. - P. scrobicu-

latum L. var. j) Flugge, Gram. Monogr., Paspalum (1810) 87. - Type: Herb. Willdenow

1574 microfiche IDC!: third from India. p.p. (B, holo,n.v.; left),

P. P. polystachyum R.Br., Prod. 1 (1810) 188; R. & S., Syst. Veg. 2 (1817) 297. - scrobicula-

tum L. var. polystachyum Stapf in Prain, Fl. Trop. Afr. 9 (1920) 576. — Lectotype:

R. Brown 6088 (K, holo), Australia, Bay of Carpentaria, 1802-1803.

P. firmum Trin., Gram. Pan. (1826) 105; Steudel, Syn. 1 (1853) 28. - P. mauritanicum Nees ex

Steudel, Syn. 1 (1853) 25, nom. superfl. (see note). - Type : Sieber Fl. Maur. 44 (LE, holo;

P, n.v.; L), Mauritius.

P. cartilagineum Presl, Rel. Haenk. 1 (1830) 216; Miq., Fl. Ind. Bat. 3 (1855) 432; Jansen, Rein-

wardtia 2 (1953) 320; Bor, Grasses (1960) 335; Henty, Bot. Bull., Lae 1 (1969) 145; Gill.,

Rev. Fl. Mai. 3 (1971) 184. - P. orbiculare Forst. f. var. cartilagineum Summerh. & Hubb.,

Kew Bull. (1930) 257. - P. cartilagineum Presl var. cartilagineum: Fosb. & Sachet, Micro-

nesica 18 (1984) 81. -Lectotype: Haenke s.n. (PR, holo), Philippines, Luzon, Sorzogon,

1792.

P. adelogaeum Steudel, Syn. 1 (1853) 27. - Type: Goering 11, 168 (P, holo), probably from

Java.

P. metzii Steud., Syn. 1 (1853) 21. - Type: Hohenacker 923 (P, holo, n.v.; L, UPS, W), India,

Nilgirris, 1851.

- P. polo F.M.Bailey, Queensl. Agric. J. 1 (1897) 234, f. l;Queensl. Fl. 6 (1902) 1813. Type

Bailey 51 (BRI, holo), Australia, Queensland, Somerset, Polo Creek, June 1897. 306 BLUMEA - VOL. 30, No. 2, 1985

P. scrobiculatum L. vai. gracillimum Domin, Bibl. Bot. 85 (1915) 288. - Type : not indicated

(?PR, holo,n.v.), Australia, Queensland, Townsville, anno 1910.

P. commersonii Lamk. var. hirsutum Jansen, Reinwardtia 2 (1953) 319. - Type: SF 4787

(Nur) (SING, holo;K), Malaya, Selangor, Kuala Lumpur, 22 Feb. 1919.

P. scrobiculatum auct. non L., p.p.: Trin., Diss. Bot. Alt. (1826) 122; Sp. Gram. Icon. 2 (1829)

t. 143 (!); Buse in Miq., PI. Jungh. 3 (1854) 382; Miq., Fl. Ind. Bat. 3 (1857) 430; Merr.,

Philip. J. Sc. 1 (1906) Suppl. 344, p.p.; Gill., Rev. Fl. Mai. 3 (1971) 185, f. 38 (!); Hsu,

Taiwania 16 (1971) 292.

P. cartilagineum Presl var. biglumaceum Fosb. & Sachet, Micronesica 18 (1984) 81. — Type :

Necker RS-292 (US, holo), Marianas, Rota I., Sabana, 1 Nov. 1945.

Differs from the typical variety by:

Culms 8-70(-130) cm long. Cauline sheaths glabrous or hairy along the margins

at the top, rarely hirsute all over. Blades 10—25(—42) cm by 4 —9(—15) mm. Pedun-

cle 0.5—1.4 mm diam. Racemes (1 or) 2—6(—14), alternate or rarely subopposite,

1.5— 1.8—2.6 1.45-2.15 5- 9(—11) cm long. Spikelets by mm. Upper glume or 7-

nerved. Sterile lemma 5- or 7-nerved, usually more or less smooth. Fertile lemma

1.6—2.3 mm long. Anthers 0.5—1.1 mm long.

Distribution. Tropical Africa (not in Mauretania, see note!), Madagascar, Mauri-

tius, Asia, to the Pacific (Mariannes, Bougainville, New Caledonia), Australia (N. Ter- ritory, Queensland, New South Wales), throughout Malesia.

Ecology. In many habitats, on many types of soil, from dry to damp, sunny to

shaded, in rural areas, on beaches and along roads. A weed in sawahs and plantations.

with A sand binder on gravel banks. Associated e.g. Fimbristylis miliacea, Imperata,

Melaleuca. Up to 1900 m altitude.

Vernacular names. Malaya: pala belang, rumput tulo santadok; Sumatra: lulan-

gan, rumput manis (Gajas), r. paitan, r. tumbag (Gajas), siansak; Bangka: gegit lalang;

2 Java: blembeman, genjoran, jampang merak (Sund.), jampang ping -basit, jukut jajampangan, tatamagan, tuton gilig; Lesser Sunda Islands: amila (Alor), semanki

batu klambu (Flores); Borneo: kumpai (illapatan), k. sikubalang, rumput rusak, r.

2 moto kaki; Moluccas: njelo (Ternate), o hoye (Halmaheira), rumput piso (Buru);

2 Celebes: cani New Guinea: fatana (Minafia, Utukap), tambuka (Kulomo), , kaleta;

wakerakampa (Waskuk).

Collector's notes. Plants pale, glaucous, leaves and sheaths midgreen, flushed

purple at base. Inflorescence light to dark green-brown. Flowers green to dark brown, odourless. Anthers dark white black. yellow to purple. Stigmas or purplish Fruit

of green. Host Cerebellapaspali.

Metcalfe of the leaf of Anatomy. (1960) gave an extensive description anatomy

two specimens identifiedfor him as P. commersonii. He observed some curious dif-

ferences between them, it might therefore just be possible that he had two different

varities in hand, e.g. var. bispicatum and var. lanceolatum.

Chromosome numbers. 2n = 20 (Tateoka, 1965).

Notes.Paspalum polystachyum was described by R.Brown (1810) in living material

seen in 'T', i.e. from the tropical coasts of Queensland and the Bay of Carpentaria. In

K there is material named by him from the Prince of Wales Island from the first area and from The first has the number '6088'and is selected the Sydney. as type. R. de Koning & M. S. M. Sosef: Malesian species of Paspalum 307

Steudel (1853) took up a Paspalum described by Nees on two Sieber exsiccates,

of Fl. Maur. 44 and Fl. Mixta 145, without realizing that the first was also the type

' P. firmum Trin. (1826) also recognized by him. He erroneously wrote mauritanicum’ thereby suggesting that the species would occur in Northwest Africa, although Nees

for the other side actually, and correctly, had written ‘mauritianum’ Mauritius on of the continent.

The type of P. cartilagineum falls within the circumscription of this variety. As explained in the introduction a taxon distinguished by an indurated fertile lemma cannot be upheld.

Fosberg & Sachet (1984) have described P. cartilagineum var. biglumaceum be- cause of the presence of a lower glume. They noted that this may also occur in the typical variety. We have also noted the occurrence of an occasional lower glume, but have not found it to correlate with an indurated fertile lemma, while it occurs in at least two of the varieties distinguished here: var. bispicatum (e.g. Monod de Froide- ville 978) and var. scrobiculatum (e.g. Monod de Froideville 1566). It must be ad-

Pacific mitted that lower glumes occur more frequently in the than elsewhere, but we

this rather think this is insufficient to distinguish a variety on haphazardly occurring feature.

Fig. 7B. Paspalum scrobiculatum L. var. horneri (Henr.) Koning & Sosef. a. Upper glume; b. sterile lemma (NGF47924).

d. horneri & - var. (Henr.) Koning Sosef, stat. & comb. nov. Fig. 7B.

P. horneri Blumea 1 f. 2. - Horner Henr., (1935) 306, Ty p e: s.n. (L, holo), Sumatra, Padang.

Differs from the typical variety by: 308 BLUMEA - VOL. 30, No. 2, 1985

Culms 15—80 cm long. Nodes glabrous or appressed hirsute to pilose. Sheaths

Blades 3—6 shortly hirsute, sometimes minutely so, rarely glabrous. mm wide,

least the usually hirsute at above, sometimes minutely so, or only along margins,

Peduncle alternate rarely glabrous. usually sparsely hairy. Racemes 1—4, or sub-

opposite when paired. Pedicels 0.2—1 mm long, usually hairy all over. Spikelets

broadly elliptic to ovate, 1.7—2.5 by 1.2—1.85 mm, orange- to dark-brown, the

Sterile lemma Fertile margins usually paler. Upper glume 5-nerved. 5- or 7-nerved.

lemma brown. Anthers c. 0.8 mm long.

Distribution. Very disjunct: Assam and Malesia: Sumatra (Padang), Philippines

(Luzon), New Guinea (East Sepik, Western and Eastern Highlands, Gulf, Central,

Morobe, Milne Bay).

Ecology. On ridges, hills, in savannah depressions and bogs, occasionally com-

mon where the natural grasslands have been cut, associated with e.g. Arundinella,

Dianella, Gleichenia, Themeda. On sand, silt, peat, 25—4000 m altitude.

Vernacular name. New Guinea: kuk (Melpa).

Collector's notes. Erect, forming strongly rooted tussocks in bogs. Leaves

green,grey. Inflorescence brown, sticky. Fruit grey.

few Note. The very disjunct distribution and the collections (mainly from Papua

hence New Guinea) would suggest a polytopic origin and an unacceptable taxon. We

cannot show one or disprove another opinion, but in view of the clear differences

with the other which than the differences between these varieties, are better (!), we

thought it advisable to retain the former species at present.

8. Paspalum urvillei Steudel — Fig. 8.

P. urvillei Steudel, Syn. 1 (1853) 24; Chase, Contr. U.S. Nat. Herb. 28, 1 (1929) 173, f. 108;

Gardner, Fl. W. Austr. 1, 1 (1952) 242; Bor, Grasses (1960) 341; Vickery, Contr. Nat. Herb.

N.S.W., Fl. Ser. 19, 1 (1961) 120; Tttrpe, Lilloa 32 (1967) 50; B.C. Stone, Micronesica 6

(1970) 217; J.W. Parham in A.C.Smith, Fl. Vit. Nov. 1 (1979) 339. — Type: Dumont-

d'Urville s.n. (P, holo), Brazil.

For further synonyms see Chase (1929).

Culms Perennials, tufted, not stoloniferous. erect, (65-)90-200(-300) cm long,

sometimes branching. Nodes glabrous, rarely with some hairs. Sheaths slightly keeled

in the upper part, the basal usually densely pilose, not reticulate, the cauline gla- brous. Ligule triangular, up to 6 mm long. Blades flat, linear, narrowed at base, (25-)

30—45(—50) cm by 5—15 mm, usually with some up to 6 mm long, white hairs at

Peduncle 1.3 the immediate base of the ligule, otherwise glabrous. c. mm diam., gla- brous. Internodes of the main axis 0.3—0.5 times as long as the subtending raceme.

Stalk c. 2.2(—4) mm long, appressed hirsute, mixed with up to 9 mm long, flattened

the lowest —13 the hairs, brown. Racemes (5 —)10 —20, alternate, (5 —)8 cm long, others gradually shorter. Rachis flat, winged, straight, frequently distally zigzag, 0.7—

Pedicels flat- 0.9(—1.1) mm wide, spikeled from the base, margins scabrous. slightly

the outer to the shorter. tened, edges ciliate, ones up 2.5(—4) mm long, inner ones

Spikelets paired, imbricate, narrowly ovate to oval, 2—2.8(—3) by 1.2—1.5 mm, R. de Koning & M. S. M. Sosef: Malesian species of Paspalum 309

Fig. 8. Paspalum urvillei Steudel. a. Upper glume; b. sterile lemma (Dissing 2527).

green-brown to brown or purplish, apex acute. Lower glume absent. Upper glume

slightly convex, as long as the spikelet, obscurely pilose, with a fringe of relatively

the long, c. 1.5 mm long, white hairs on the marginal nerves framing spikelet, 3- or

5-nerved, concolorous with to slightly darker than the intervenium, midnerve promi-

Sterile flat, not wrinkled, the Fertile lemma nent. lemma as upper glume. slightly

convex, 3-nerved, thickly cartilaginous, glabrous, minutely papillose-striate, pale

green. Palea flat to slightly convex, 2-nerved, as the lemma. Anthers slender, 0.8—1

dark round mm long. Stigmas brown to purple (i. s.). Caryopsis to round-elliptic,

plano-convex, c. 1.5 mm long.

Distribution. Native to northern South America, introduced in many (sub)tropi-

cal areas, escaping here and therein Malesia: Malaya, Java, Philippines (Luzon), New

Guinea (Biak, Bismarck Archipelago).

Ecology. Along ditches and roadsides, in waste areas, usually on rather moist

soil, up to 2600 m altitude in Luzon.

Vernacular name. English: vasey grass.

Chromosome number. 2n = 40 (e.g. Tiirpe, 1967).

fodder Uses. Cultivated as a grass, readily eaten when young, but becoming tough

and unpalatable with age (Bor, 1960). Sometimes cut for hay (Chase, 1929).

Note. The above description is based on Malesian material, usually cultivated;

South American material was compared where necessary.

Van Leeuwen BIAK 6 is somewhat aberrant the of by presence only 5 racemes.

9. Paspalum vaginatum Sw. — Fig. 9.

P. vaginatum Sw., Prod. (1788) 21; Miq., Fl. Ind. Bat. 3 (1857) 433; Camus, Fl. Gen. I.-C. 7

(1922) 394; Blatter & McCann, J. Bombay Nat. Hist. Soc. 32 (1928) 640; Chase, Contr.

U.S. Nat. Herb. 28, 1 (1929) 41, f. 19; Burk., Diet. Econ. Prod. 2 (1935) 1675; Backer 310 BLUMEA - VOL. 30, No. 2, 1985

in Heyne, Nutt. PI. Indon. ed. 3, 1 (1950) 202; Gardner, Fl. W. Austr. 1, 1 (1952) 241;

Schmid, l'Agron. Trop. 13 (1958) 304, f. 48-2; Bor, Grasses (1960) 341; Vickery, Contr.

Nat. Herb. N.S.W., Fl. Ser. 19, 1 (1961) 117; Monod de Froid. in Backer & Bakh.f., Fl.

Java 3 (1968) 569; Debray, Bull. Centre Etud. Rech. Sc., Biarritz 7 (1969) 585; Henty, Bot.

Bull., Lae 1 (1969) 146; Gill., Rev. Fl. Mai. 3 (1971) 182, pi. 24c; Hsu, Taiwania 16 (1971)

294; Kerguelen, Lejeunia n.s. 75 (1975) 223; Sindhe, Taxon 24 (1975) 368; Taxon 26

(1977) 268; Hsu, Fl. Taiwan 5 (1978) 588. - P. distichum L. var. vaginatumSw. ex Griseb.,

Fl. Brit. W. Ind. (1864) 541. — Digitaria vaginata Magnier, Scrinia 6 (1887) 120. - Sanguina-

4 ria vaginata Bubani, Fl. Pyr. (1901) 258. -Type: Swartz s.n. (S, holo), Jamaica.

P. littorale R. Br., Prod. 1 (1810) 188; R. & S., Syst. Veg. 2 (1817) 316; Trin., Sp. Gram. Icon. 1

(1828) t. 112. - P. vaginatum Sw. var. Trin., Diss. Bot. Alt. (1826) 95. - P. vaginatum Sw.

var. littorale Trin. ex Buse in Miq., PI. Jungh. 3 (1854) 383; Miq., Fl. Ind. Bat. 3 (1857) 433.

-P. distichum L. var. littorale F.M.Bailey, Queensl. Gr. (1888) 23.- Lectotype: R. Brown

6091 (BM, holo;K), Australia, Queensland, Cumberland Islands, Prince of Wales I., 1802.

P. boryanum Presl, Rel. Haenk. 1 (1830) 209; Miq., Fl. Ind. Bat. 3 (1857) 432. - Type : Haenke

s.n. (PR, holo), Philippines, Luzon, Sorzogon, 1792.

Rottboellia ? uniflora Cunningh. in Hook., Comp. Bot. Mag. 2 (1837) 371. - Type: Cunning-

ham s.n. (K, holo,n.v.), New Zealand, North Island, 1834.

P. distichum auct. non L.: Bor in Rechinger, Fl. Iran. 70 (1970) 494; Vickery, Fl. New South

Wales no. 19 (1975) 135; Simon, Techn. Bull. Bot. Br., Brisbane 4 (1980) 72; Wheeler et al..

Grasses New South Wales (1982) 229.

For further Chase non-Malesian listed her have synonyms see (1929) (some synonyms not by

here been included).

Perennials, not tufted, stoloniferous. Stolons leafy, rooting at the nodes. Culms

ascending to erect, (15—)25—50(—100) cm long, rarely branching. Nodes glabrous.

Sheaths not keeled, glabrous, the basal ones not reticulate. Ligule collar-shaped, 0.5—

1.1 mm long. Blades strongly distichous, flat or involute, linear, narrowed to the

with base, (2 —)8—18 cm by (1 —)1.5—4(—6) mm, usually some c. 0.5 mm long,

white hairs at the immediate base of the

ligule, otherwise glabrous. Peduncle c. 1.1

mm diam., Stalk glabrous. up to 1.5 mm

long, frequently with some c. 1 mm long,

white hairs, pale to brown. Racemes 2,

opposite, rarely up to 5, then the upper 2

opposite, the lower ones alternate, 2—5

(—8) cm long. Rachis flat, sometimes

more or less grooved, straight to slightly

zigzag, 1—2 mm wide, without spike lets

immediately above the stalk, naked for

the lower c. 5 mm, margins smooth. Pedi-

cels 2-edged, 0.3—0.8 mm long, edges

finely ciliate, sometimes obscurely so.

Spikelets solitary, at least the upper ones

3—4.5 1.3—1.6 imbricate, oblong, by mm,

glabrous, pale green, apex acute. Lower

glume rarely developed (e.g. in the type Fig. 9. Paspalum vaginatum Sw. a. Upper of P. then glume; b. sterile lemma (Backer 20700). vaginatum), a minute, oblong R. de Koning & M. S. M. Sosef: Malesian species of Paspalum 311

scale. Upper glume slightly convex, as long as the spikelet, glabrous, nerves 3—7, con-

colorous with the intervenium midnerve often obscure. Sterile lemma flat, often

weakly, transversally undulate in the lower half, 3-, or 5-, or 7-nerved, midnerve

the 3- 5-nerved, prominent, as upper glume. Fertile lemma convex, (0- or) or perga-

mentaceous, smooth, pale green, the apex with some short, stiff cilia, otherwise gla- brous. Palea flat, 0- or 2-nerved, as the lemma. Anthers slender, 1.2—1.5 mm long.

Stigmas dark brown (i.s.). Caryopsis ovate, plano-convex, c. 2.5 mm long.

coasts of Malesia: Distribution. Sea (sub)tropical areas, in Malaya (Pinang,

Perak, Pahang, Selangor, Johore), Singapore, Sumatra (East Coast), Riau (P. Bintan),

Java (throughout, also in lowland, inland saline areas, fide Backer, 1950; see note),

Kangean Islands, Lesser Sunda Islands (Bah, Sumbawa), Borneo (Sarawak, Brunei,

Sabah), Philippines (Luzon), Celebes (Kendari), Moluccas (Halmaheira, Ceram,

Tenimbar), Aru Islands (Dobo), New Guinea (Vogelkop, Adi I., Gulf, Morobe, Mus-

sau I.).

Ecology. Halophilous, favouring exposed places, rarely in the shadow, locally

vegetation forming: on beaches, along tidal pools, at river mouths, mangrove margins,

also of and salt swamps, can withstand flooding, on quays, in plantations coconuts

low altitude sawahs and inland saline areas at least in Java. Associated with Casuari-

Ischaemum at na, Cyperaceae, Ipomoea pes-caprae, spp., Spinifex. Usually sea level,

50 altitude salt rarely up to m in ponds.

Vernacular salt Java: names. English: water paspalum; Malaya: rumput dawai; asianan.

stem Collector's notes. Mat-forming with long runners, main .and bases of

secondary ones red, leaves slightly fleshy. Spikelets light green. Anthers dark purple, black. Stigmas black.

Chromosome numbers. 2n = 20 (e.g. Hsu, 1971), 40 (e.g. Sindhe, 1977), 60

(e.g. Sindhe, 1975).

Uses. Readily eaten by cattle and possibly of some nutritional value as a fodder

useful for on swampy, saline soils. Not very hay as it tends to dry slowly. An impor-

tant soil binder (Burkill, 1935;Backer, 1950).

Notes. For the nomenclatural problems that have involved the name accepted

here see under P. distichum.

One culm of a collection from Adi I., New Guinea {BW 6336, Kalkman, L) has

two racemes each with a secondary branch, a very rare abnormality.

The species has been mentioned by Backer (1950) for inland saline areas in Java.

A few inland collections were available, but they were always from areas only a few kilometres from the coast, e.g. Backer 23434 (BO) ('W. of Batavia, moist grassland,

not salt, very unusual habitat for the species'), 24533 (BO) ('Pradjekan, Pasoeroean,

sandy bank of kali, abnormal locality, the grass usually grows on silty clay'), several

from inland collections from Bangil, which may be stations. Qason Laarman G-60

of (BO) is from 20 m altitude near Pandar, S. Panarukan, Besuki. Elmer 18140 (K, L)

from the label further information. is Mt Maquiling, Luzon, but carries no The spe-

cies is not mentioned by Van Steenis in his survey of coastal plants occurring near

inland salt wells in Java (in Backer & Bakh. f., Fl. Java 2, 1965, '53'). - 312 BLUMEA VOL. 30, No. 2, 1985

APPENDIX

Paspalum scrobiculatum L. var. lanceolatumKoning & Sosef, var. nov.

P. lamprocaryon K. Schum. in Engler, Pfl. W. 0. Afr. C (1895) 100. - Type : Stuhlmann 3901

(B, holo,n.v.), Tanzania, Bukoba.

P. auriculatum auct. afr. non Presl: see Clayton & Renvoize, Fl. Trop. E. Afr., Gram. 3 (1982)

609 for references.

verumtamen Taxon cum Paspalo auriculato Presl (vero) confusum, nodis a vaginis involutis,

laminibus linearo-lanceolatis, 7-21 cm longis, 12-27 mm latis, basi rotundatis, racemis 2-4,

spiculis 2-2.7 mm longis bene distinctum. - Typus: Stolz 1413 (L, holo), Malawi, Nyassa

Highlands, Kyimbila Station, 1100-1400 m alt., 3 July 1912.

Confused with the true Paspalum auriculatum Presl, however different from that

by the sheaths enveloping the nodes, blades linear-lanceolate, 7—21 cm by 12—27 mm, rounded at base, racemes 2—4, spikelets 2—2.7 mm long.

Distribution. Tropical Africa, South to Angola, Zimbabwe.

Ecology. Damp places and forest margins, 50-2000 m altitude.

Note. of the had indurated sterile which the Most specimens seen lemmas, in other and then. be this form. varieties occur only now It may more common in

ALIEN AND EXCLUDED SPECIES

A number of species have been found once or twice apparently escaped from botanical have been introduced the gardens, experimental stations, or recently in

These have included the whether area. been in key, only, as it cannot yet be estimated they will really maintain themselves.

at another in The many Malesian species one time or placed Paspalum, but in- cluded in other genera, at presently have not been taken up here, as that information seemed to be a waste of paper.

Paspalum decumbens Sw., Prod. (1788) 22; Chase, Contr. U.S. Nat. Herb. 28, 1

(1929) 92, f. 48; Jansen, Reinwardtia 2 (1953) 322; Monod de Froid. in Backer &

Bakh.f., Fl. Java 3 (1968) 569. — Type : Herb. Swartz (S, holo; B, M, P, n.v.),

Jamaica.

This South American species has been introduced in West Java and may have escaped from the Botanical Gardens in Bogor, although it was never cultivated there on purpose. It was first found in 1949 (Monod de Froideville 979, BO).

Paspalum notatum Flugge, Gram. Monogr., Paspalum (1810) 106;Chase, Contr. U.S.

Nat. Herb. 28, 1 (1929) 64, f. 32; Parodi, Rev. Argent. Agron. 15 (1948) 54; De

Castro, Garcia de Orta 12 (1964) 58; Gould, Grasses Texas (1975) 506; Madulid,

Acta Manil. A, 14 (1975) 121; L.B. Smith et al., Fl. II. Catarinense, Gram. (1982) R. de Koning & M. S. M. Sosef: Malesian species ofPaspalum 313

944, f. 190a-d. - Lectotype: Ventenat s.n. (B, KIEL, MO, n.v.), St. Thomas,

1802.

This species has been introduced recently in formerly Portuguese Timor (Cinatti

111, LISC, n.v.), Luzon (fide Madulid, 1975), and at least in Flores (Veldkamp 6998,

BO, L, Herb. Verheijen in Ruteng) as a grass for lawns and pastures. It grows well on

low-fertility soils probably due to symbiosis with Azotobacter paspali, a nitrogen

fixing bacterium (Dobereiner, 1966, cited by Smith et al., 1982). It is very resistant

to drought and spreads rapidly and aggressively by its rhizomes and has therefore

been used for soil conservation and the prevention of erosion in Uganda and the

U.S., where it is known as Bahia- or Pensacola Bahia grass; in Timor it was called

fodder. 'rumput pencasilan' (sic) and was consideredto be a good

next to the which Generally two varieties have been distinguished typical one,

according to Parodi (1948) would differ by somewhat larger spikelets and longer

racemes. The collection from Flores belongs to var. notatum.

Paspalum plicatulum Michx., Fl. Bor. Amer. 1 (1803) 45; Chase, Contr. U.S. Nat.

Herb. 29, 1 (1929) 214, f. 129; Hill, Papua New Guinea Agric. J. 22 (1970) 44;

— Gallash, Papua New Guinea Agric. J. 22 (1971) 63. Type : Herb. Michaux s.n.

(P, holo, n.v.), U.S.A., 'Georgia, Florida'.

Two cultivars have been tried in Papua New Guinea (cv. Rodds Bay and cv. Hart-

ley), which seed freely. They are highly palatable to cattle. As they are good pioneers

in of low where and grow well areas reasonably fertility, they are bound to escape

introduced collection 41935 from (Hill, 1970). Only one was seen (NGF Henty \ L)

a cultivation plot in Bubia.

Yield tests reported by Gallash (1971) show fairly high production figures, higher

even than those of the for grazing purposes more commonly used P. conjugatum.

Paspalum tenellum Willd., Enum. PI. (1809) 89; Chase, Contr. U.S. Nat. Herb. 28, 1

(1929) 112, f. 62. — P. elegans Flugge, Gram. Monogr., Paspalum (1810) 183;

F.-Vill., Nov. App. (1882) 310, nom. superfl. — Type : Herb. Willdenow 1608

(B, holo, n.v.; microfiche IDC!, lefthand specimen), cult, in Berlin.

A number of other species have been mentioned by F.-Villar (1882), some have

never been found again and no records exist, so they again may be examples of the

unreliability of this work. (See for comment also Merrill, Sp. Blanc., 1918, 16-17.)

The species has therefore not been included in the key.

Paspalum virgatum L., Syst. Nat. ed. 10, 2 (1759) 855; Hitchc., Contr. U.S. Nat.

Herb. 12 (1908) 116; Chase, Contr. U.S. Nat. Herb. 28, 1 (1929) 197, f. 119;

1846. — P. Herb. Baum, Canad. J. Bot. 45 (1967) Type : Browne s.n. in Linne

80.26 microfiche Jamaica, no. (LINN, n.v., IDC), see note.

PI. 3 384. Paspalum sp. : Buse in Miq., Jungh. (1854) 314 BLUMEA - VOL. 30, No. 2, 1985

Once collected by Junghuhn in Java, probably, as Buse also suspected, cultivated.

It has not been found again.

Contrary to Baum (1967) P. virgatum was not based 'on one element only', i.e.

the description and plate of Sloane and hence on Sloane's specimen in BM, which

has the but which Baum appointed as type, is most unlikely to have been studied or

The latter of his which had even seen by Linne. gave a diagnosis own to be based on actual material, e.g. a P.Browne specimen (LINN), which he as usual did not cite, and which he matched with Sloane's work. See Steam's introduction to the facsimile edition of the ed. 1 and for similar of Species Plantarum, (1957, p. 108,113) a case a mistaken typification Veldkamp (Taxon 33, 1984, 95). In any case, the name has been lectotypified previously by Hitchcock (1908), which was followed by Chase

(1929) and this has to be followed.

Paspalum wettsteinii Hackel, Denkschr. Wiss. Math. Naturw. Akad. Wien 79 (1906) 5

(preprint), (1908) 66; L.B. Smith et al., Fl. II. Catarinense, Gram. (1982) 994, in

syn. - Type : Wettstein & Schiffners.n. (?W, holo, n.v.), Brazil, Sao Paulo, Con-

ceicao, Rio Branco, July 1901 (not received from W).

Paspalum wettsteinii has been cultivated in Australia, Two collections from Papua

New Guinea have been seen, LAE 72440 Henty (L) and Sterly 1658 (L). The first was cultivated in Abunaka, Morobe Dist., the second occurred along a roadside near

Noglkanmambuno, Gembogl subdist., Chimbu Dist. Sterly noted that it had been cultivated and recorded the pidgin remark 'namba wan' (= + 'excellent'), which indi- cates the enthusiasm with which this cultigen has been received there.

Smith et al. have placed this name in the synonymy of P. conspersum Schrad.

of Unfortunately the type P. wettsteinii was not received from W, so we could not check whether this was correct. Our specimens, however, matched Hackel's descrip- tion from from material and these and of P. conspersum we conclude that the two

differs are distinct taxa. The present species clearly from the latter by e.g. the nar- rower rachis and glabrous spikelets.

Index of collectors

Only numbered collections have been included. Specimens cited in literature but not seen have been included with their identities between brackets when these seemed acceptable, other-

been wise they have omitted. Paspalum conjugatum is underrepresented, because it was specifical-

asked include this in the it is and without ly not to species loans, as very common any special problems. The numbers correspond with the species as presented above, the letters with the initials ofthe alien species (which were included in the key and the short survey above).

Adj. Landbouwer Pamekasan 10: 7a - Adj. Veearts Gorontalo 32: 4 - Aet 313: 4; 796: 4 -

Aet & Idjan 735: 1 - Afd. Binnenvisscherij 32: 9 - Agric. Res. Centre Tuaran 563: 2 -

Agric. Stat. Tuaran 528: 4 - d'Albertis 257: (4) - Allers 184-81: 1 - Alston 13068: 7c;

16510: 7c; 17046: 7c - Alvins 2166: 4 - Amdjah 111: 4; 811: 7c; 837: 4 — Anang 278: 4;

456: 1; 503: 7c; 584:1 - Anonymus 8045: 7a - Anta 676: 7c; 887a: 1 - ANU 514 (Walker):

5; 1742 (Kellman): 1; 6189 (Wheeler): 7d; Asdat 183: 4 Atasrip 157: 7c. R. de Koning & M. S. M. Sosef: Malesian species ofPaspalum 315

Backer 54: 4; 197: 7c; 860: 4; 937: 1; 1023: 7c; 1375: 1; 1834: 4; 1919: 1; 2096: 4; 2392: 7c;

2578: 4; 3393: 7c; 4127: 4; 4229: 4; 5137: 9; 5148: 7c; 5154: 9; 5474: 7c; 5537: 5; 5817:

4; 6198: 7c; 6658: 7a; 6857: 7b; 7134: 7c; 7319: 1; 7573: 9; 7670: 1; 7817: 1; 8132: 7a;

8257: 9; 8434: 9; 8746: 5; 8747: 1; 9165: 1; 9969: 4; 10050: 7c; 10052: 1; 10560: 4;

10565: 7c; 10744: 2; 11791: 9; 11887: 1; 11892: 4; 12123: 4/7c; 12941: 9; 13997: 1;

14079: 4; 14276: 7c; 14541: 7c; 14594: 5; 14814: 7c; 14924: 4; 15245: 9; 15419: 1;

15599: 1; 15668: 5; 15881: 7c; 16309: 9; 17109: 7c; 17262: 4; 17421: 1; 17430: 7c;

17651: 9; 18280: 7c; 18477: 7c; 18578: 4; 18999: 1; 19112: 9; 19115: 7a; 19547: 9;

20043: 7c; 20700: 9; 21173: 9; 21389: 9; 22445: 4; 23434: 9; 23532: 1; 23539: 4; 23540:

7c; 23770: 7b; 24093: 7c; 24143: 9; 24507: 9; 24533: 9; 24689: 9; 25077: 2; 26579: 7a;

27105: 7a; 27676: 9; 27730: 7a; 28026: 9; 28602: 9; 29143: 7c; 29251: 9; 29701: 9;

30125: 1; 30327: 7c; 30991: 9; 31073: 9; 31831: 7c; 31835: 4; 31837: 4; 31850: 9; 36053:

9; 37556: 9 - Bakhuizen van den Brink 9: 7c; 44: 1; 70a: 4; 141: 1; 180: 4; 707: 2; 1086:

7c; 1669: 7c; 2018: 9; 2022: 9; 2767: 4; 2768: 7c; 4446: 4;4546: 4; 7796: 7c; 14446: 4 - - Barber 83: 9; 300: 4; 341: 9 bb 9013 (Ostwald): 7c; 11158 (Japing): 7c - Beccari B-110:

4; B-3528: 9 - Beguin 15: 7c; 48: 1; 60: 2; 67: 7c; 180: 1; 614: 7c; 2050: 7c; 2278: 9 -

Beumee 10a: 4; 782: 7a; 855: 7c; 1066: 1; 1115: 7c; 1155: 7c; 1176: 7c; 1555: 7a; 1703:

7c; 1747: 7c; 2208: 7c; 2840: 7c; 2892: 7c; 3172: 4; 3609: 7c; 3717: 7c; 3772: 7a;4156: - 7c; 4811: 4; 4835: 7c; 5559: 7c - Bloembergen3010: 1; 3516: 4;4524: 7b Boerlage667:

7c - Bor S-14: 5; S-18: 7c; S-30: 4 - Brass 521: 4; 1229: 9; 3935: 1; 5922: 4; 6285: 9;

6297: 7c; 6341: 4; 6948: 7c; 7529: 7c; 8302: 7c; 8325: 1; 13784: 7c; 21674: 1; 22037: 7c; - 22080: 9; 23815: 7c; 24420: 6; 25116: 6; 28208: 6 - vanBreemen 21: 7c; 31: 2 Brooke

8268: 5; 9833: 5 - D.A.L.I. Brown 10: 7c; 11: 7c - BS 318a (Shan): 7c; 843 (Foxworthy):

(7b); 941 (Mangubat): 4; 1449 (Ramos): 4; 1653 (Robinson): 1; 1956 (Ramos): 4; 2454

(Native coll.): 4; 9323 (Merrill): 5; 10083: (7c); 10979 (Ramos): 4; 10984 (id.): 5; 11621

(Robinson): 4; 11667 (id.): 4; 14512: (4); 16878 (Servinas): 1; 18908 (McGregor): 1;21684

(Ramos): 4; 24435 (id.): 7c; 26002: (4); 26675 (Fenix): 5; 27378 (Ramos): 7c; 29504

(Ramos & Edano): 4; 31512 (id.): 7c; 31758 (Santos): 5; 32275: (7c); 35186 (Farinas): 1;

43974 (Ramos & Edano): 7c;43975 (id.): 4; 44275 (id.): 4 - Biinnemeijer 144: 7c; 193: 7c;

218: 1; 791: 5; 1060: 1; 1320: 1; 1323: 5; 1386-b: 5; 1394: 1; 1570: 5; 1638: 7c; 1717: 5;

1854: 5; 2158: 7c; 2246: 1; 2807: 1; 2809: 5; 2911: 7c;3549: 1;3722: 7c;4248: 1;4322:

7c; 4789: 1; 4791: 5; 5597: 1; 5603: 5; 5998: 4; 6005: 1; 6220: 1; 6221: 5/7c; 6258: 5;

6270: 4; 6277: 1;6333: 7c; 6575: 4; 6617: 5; 6910: 1; 7015: 5; 7216: 4; 7398: 1;7400:4;

7403: 7c; 7501: 5; 7618: 5; 8011: 7c; 8171: 1; 8282: 5; 10744: 1; 11022: 5; 11028: 7c;

11142: 7c; 11719: 1 - Burkill & Haniff 464: 9 - Burkill& Shah 945: 7c - Buwalda 2729: 1;

2788: 7c; 3394: 1;3744: 1;4532: 7c;4613: 9;4851: 9;4885: 1;5798: 1;6047: 9; 7875: 1;

7911: 4; 8160: 7c - Buijsman 119: 2 - BW 723 (Versteegh): 1;6283 (Kalkman): 1; 6336

(id.): 9; 8366 (Versteegh & Vink): 1.

Carr 11025: 1; 12957: 6; 12972: 6; 13598: 6; 14282: 6 - CF 20384 (Strugnell): 7c - Cheesman - 220: 2 - Cinatti 11a: 5; lib: 2; 51: 7c; 111: n; 116: 7c- Clason A20: 1 Clason-Laarman

= G-20: 9 - Clemens 399: (4); 17953: 7c; 17954: 9; 17955: 9; 18235: 7c; 30264: 7c; 32616

- - 34124: 1 - Coert 72: 7c; 728: 1; 791: 9; 831: 9; 1728: 7c Coode 3773: 6; 3782: 7c

Copeland 371: 4; 575: (1) - Craven & Schodde 717: 7c - Cruttwell 169: 7c; 1609: 5; 1616:

7c - Cuming 720: 4/7c; 2410: 4 - Curtis 492: 5;493: 1; 1957: 9.

van Daalen 304: 1; 419: 4 - Dakat 2: 5 - Dalenberg 155: 7c - Dames 84: 1 - Danser 5386:

7c; 6023: 2; 6453: 7c; 6459: 4; 6795: 7c; 6866: 7c - Darbyshire 185: 6; 827: 5 - Darby-

shire & Hoogland 7864: 1 - DeVore & Hover 101: (4); 180: (1) - Dihm 109: 1; 121: 4 -

Dilmy 1406: 9 - Dissing 2527: 8 - Djadoek 1113: 7c — Docters van Leeuwen 9252: 1 -

Docters van Leeuwen-Reijnvaan 3765: 9; 5552: 5 - Dorgelo 168: 9; 637: 7a; 694: 7c;

3243: 7c.

- Elbert 4254: 7b - Elmer 5759: 5; 16494:1; 16727:4; 18140: 9; 18184:1 Elsener H-213:5 -

Endert 1570: 1; 1679: 1; 1746: 7c; 1895: 7c; 2096: 7c - Enoh 95: 7c; 334: 7c — Everaarts

74: 1. 316 BLUMEA - VOL. 30, No. 2, 1985

- - Fairchild 178: 1 - Farinas 115: 7c; 116: 4; 117: 3 Feuilletau-de Bruyn 205: 7c; 342: 1 Fitz-

gerald 117: 7c - Flenley 61: 7c - FMS 14470 (Symington): 4; 37620 (id.): 4 - Forbes

J-1266: 7b; T-3462: 7c; T-3472: 7b - Forster F-10: 5; F-74: 4; F-84: 7c; F-137: 7c; F-144:

4; F-147: 7c - Foxworthy 843: (7b) - Franch 173: 1; 174: 4.

Gezagh. Manggerai 2: 5 - Gianno 110: 7c; 187: 1 - Gibbs 2773: 7c - Gilliland 5179: 7c;5231: - 5 - Gisius 76: 2 Gjellerup 95: 1 — Goering 11-168: 7c — Gouv. Veearts Watampone 13: 7c;

25: 7a - Grimes 1100: 7c - Groot 9: 7c - Guggitz 36: 7c.

- Hallier f. 103: 7c;631a: 7c;632b: 4;635a: 5;663: 1;C-156: 7c Hamel 1100: 4 - Harmsen6:

7c; 27: 7c - Harreveld-Lako 27: 7c -'t Hart & van Leeuwen K-15: 7c; M-10: 4; M-35: 4;

MA-12: 7c; MA-14: 1; MA-18: 7c - Hartley 11609: 7c; 11600: 1 - Hasskarl 647: 7c -

- Hellwig 407: 7c Heyligers 1340: 7d — Hiepko & Schultze-Motel 527: 9; 530: 6; 605: 4;

1018: 1 - Hochreutiner 1375: 7c; 2113: 5 — Hollrung 828: 4 - Hommel051e: 7c;078d: 9;

223c: 7c - Hoogerwerf 4: 7c; 227: 7c - Hoogland 3289: 6; 3368: 1; 3476: 6; 3681: 1;

4628: 1; 4640: 7d; 4806: 7c; 9012: 1 - Hoogland & Craven 10134: 7c; 10142: 4 - Horner - 1: 7c - Horsfield 962: 7c - Hose 4: 7c; 26: 4 - Houtvester Besoeki 14: 1 Houwing 110: - 7c; 192: 7c; 269: 7c; 860: 7a; 1010: 7c Hullett482: 5 - Hume 7266: 7c; 7337: 4; 7436:

4; 7477: 7c; 7511: 4.

Iboet 273: 1;334: 7c - Idjan & Mohtar 58: 7c; 119: 1 - Iwatsuki et al. S-148: 5; S-449: 1

Jaag 263: 7b; 1164: 7c; 1186: 7c - Jacobs 4985: 4; 5579: 1 - Jacobson 117: 5 - Janaki Ammal

10: 4 - A.H. Jansen 17: 3 - H. Jensen 222: 9 - Jermy 4417: 7c - Jochems 313b: 4; 1680:

- 7c; 3131: 7c; 3305: 7c J.K. de Jong 22: 7c.

Kalkman 4171: 1 - Karta 187: 5; 368: 7c — (Kuswata) Kartawinata 289: 9; 1373: 3; 1425 : 4 -

Kasik 54: 4; 57: 1;76: 1; 107: 5 - Kasim b. Rajab 544: 2 - Kaudern 114: 7c; 253: 1;462: 4

- -Kelly 25: 1 - Kern 7274: 1 Kievits 50: 7c; 671: 7c; 1483: 7c; 1628: 7c; 2159: 7c; 2186:

7c; 2190: 7a; 2469: 7c; 2655: 7c; 2807: 7c; 3022: 7a;3111: 7c;3318: 7c - Kingston 34: 6

- Kjellberg 377: 9; 1182: 4 - Kleinhoonte 324: 4 - Kneucker 607 (Merrill): 1; 803 (Merrill

& McGregor): 7c; 804 (Merrill): 2; 805 (id.): 9 - Ktfe & Olsen 1728: 9 - Kooper497: 9 -

Koorders 17272: 7c; 17274: 1; 17275: 1; 22308: 1; 23079: 1; 23503: 1; 23881: 1; 26208:

7c; 27171: 1; 27727: 7c; 35284: 7a; 41073: 7c; 41345: 7c; 41408: 1; 42216: 5; 42421: 1;

- 42514: 7a; 44229: 4 - Kooy 695: 1; 698: 7a; 765; 1; 771: 4 - Kornassi 456: 1; 766: 1

- Kostermans 2849: 1; 18161b: 7c; 21675a: 7c - Kostermans & Anta 357: 5 Kostermans &

- - Soegeng 130: 6 - Kostermans & Wirawan 428: 4 Kostermans & van Woerden 23: 1

- Kramer 165: 1 - van Kregten 50: 7c; 56: 1 - Kreulen 29: 1 - Kulescha 18: 7c; 44: 9 Kurz

120: 7c; 3023: 5 - Kuswata, see Kartawinata.

LAE 50291 (Stevens): 1; 50292 (id.): 7c; 53708 (Stone & Streimann): 1; 57130 (Andrew): 5;

64022 (Howcroft): 7d; 66580 (Barker & Vinas): 7c; 67671 (Barker): 1; 72440 (Henty); w; - 72458 (id.): 6 - Lam 2618: 4 - Landbouwconsulent G: 4; J: 7c Lauterbach 647: 4 -

Leefmans 161a: 1; 161b: 1; 161c: 1; 161d: 1 - van Leeuwen 3: 7c; BIAK-6: 8; DI1-6: 7c;

KAB-3: 7c; KAB-6: 7c; POE-4: 7c - Lobb 131: 7a; 139: 7c - Loenen 10: 7c - Loher 1738:

7c; 1740: 7c; 1741: 7c; 1742: 4; 1743: 4; 1744: 4; 1746: 9; 1747: 9; 1748: 9; 1749: 9;

1750: 7c; 1751: 7c; 5035: 7c - Loogen7: 7c - Lorzing 120: 7c; 387: 1;3068: 7c; 3114: 7c;

3226: 9; 3437: 7c; 3450: 4; 3486: 9; 3744: 4; 4963: 7c; 4979: 7c; 6466: 5/7c; 6644: 4;

7280: 9; 7902: 7c; 8562: 5; 8681: 7c; 9090: 9; 9242: 7c; 9789: 7c; 9521: 4; 9631: 4;

11073: 4/7c; 12502: 1; 12673: 7c; 12914: 4; 12992: 1; 13123: 1; 13141: 9; 13249: 4/7c;

13296: 4; 13588: 5/7c; 15464: 5 - Lorzing & Jochems 7560: 7c - Lutjeharms 3828: 7c;

5091: 4; 5238: 1.

McGregor 61: (1); 1250: (4) - McKee 1709: 4 - Main 413: 1;679: 1; 1741a: 7c; 1741b: 5;

1742: 5; 1988: 7c; 2052: 1; 2073b: 1 - Marche350: 7c; 416: 5 - Maxwell 81-67: 9 - Mayr

88; 1 - van der Meer 917: 7c; 918: 7c - van der Meer & van den Hoed 2002: 9 — Merrill 8:

7c; 34: (1); 140: (4); 140a: (4); 478: (4); 820: 7b; 1137: (1);4253: 7c;4793: 5; 5229: (9);

5361: (7b); 5455: 4; 7116: 9; 7292: 7c; Philip. PL 158: (7b); 576: (7c); 1956: (4); Sp. Blanc.

1035: 7c - Meijer 466: 7c; 1853: 4; 2021a: 1; 5875: 7c; 9640: 7c - Meijer-Drees 522: 1 -

Mondi 47: 7c - Monod de Froideville 623: 1; 672: 1; 913: 7c; 916: 5; 931: 9; 949: 5; 978: R. de Koning & M. S. M. Sosef: Malesian species of Paspalum 317

7c; 979: d; 1054: 4; 1141: 7a; 1190: 4; 1204: 7c; 1210: 7c; 1288: 7c; 1354: 7c; 1432: 7a;

1518: 5; 1566: 7c; 1607: 4; 1761: 4; 1808: 4; 1827: 4; 1831: 7c; 1858: 7c; 1872: 7c; 1881:

4; 1911: 7c; 1919: 7a - Morley et al. 29: 1 - Motley 43: 7c; 83: 9; 127: 7c; 133: 7c; 134: 9;

998: 4 - Mousset 20: 7c; 69: 1; 1047: 2 - Murata et al. B-25: 4;B-30: 1; B-115: 7c;B-121

4; B-312: 7c;B-558: l;B-579: l;B-3420: 4: B4576: 5. - Native coll. 5772: 7c Nedi: 380: 1; 404: 7c - Nedi & Idjan 125: 7c; 150: 7c - New Guinea

Dept. Agric. H-2868: 5 - NGBF 1030 (Argent): 1 - NGF 3162 (no coll.): 6; 3601 (Fryar):

1; 3610 (id.): 7c; 3612 (id.): 7c; 3629 (id.): 7d; 3878 (Womersley): 1; 9874 (Henty): 9; 9875

(id.): 7c; 10427 (Grey & White): 7c; 15700 (van Royen & Millar): 1; 16874 (Henty): 7d;

17410 (Epstein): 1; 22002 (Gillison): 4; 29526 (Coode): 7c; 29631 (Coode et al.): 1; 34699

(Croft & Lelean): 5; 35182 (Millar & Dockrill): 1; 38634 (Henty & Katik): 7c; 38851 (Henty

& Streimann): 7d; 40383 (Coode): 1; 41316 (Croft & Islands): 1; 41341 (Croft): 1; 41935

43414 44005 (Henty): p; (Mann & Vandenberg): 1; (Kairo & Streimann): 6;44218 (id.): 6; - 47924 (Kairo): 7d; 49125 (Henty & Barlow): 7c - Nooteboom 1190: 1 Nyman 148: 7c;

160: 7c; 1119: 7c.

Oersipuny 20: 7c - Ohwi45: 7c - Olsen 795: 7c - van Ooststroom 12512: 4; 12987: 1; 13587:

9; 13625: 1 - Ophof & de Wit 4057: 1 - Ottolander 415: 7c. - Peekel 41: 5 - Pleyte 411: 1; 490: 7c; 909: 5; 1142: 1 PNH 10300 (Mendoza & Convocar):

7c; 11296 (Edano): 7c; 16567 (Soriano): 7c; 16864 (Sulit & Conklin): 1; 17062 (Sulit): 4;

18097 (Farinas): 7c; 19086 (Conklin): 4; 20377 (Mendoza): 5; 20457 (id.): 7d; 33167 (id.):

3; 33233 (Steiner): 7c; 33670 (Gachalian): 7c; 34386 (Edano): 7c; 34767 (Steiner): 1; 35186

(Farinas): 1/4; 35189 (id.): 4; 36242 (Mendoza): 4; 37191 (Edano): 1; 38537 (Edano &

Gutierrez): 1; 39074 (Edano): 4; 42137 (Mendoza): 1; 42559 (id.): 7c; 76715 (Conklin et

al.): 1; 76793 (Mendoza & Buwaya): 7c; 78165 (Gutierrez): 1; 80661 (Conklin & Buwaya):

3; 97738 (Mendoza): 4; 117291 (Gutierrez): 7c; 117984 (Madulidet al!): 4; 118126 (id.): 4;

118203 (id.): 7c - Polak 286: 5; 721: 5 - Poore 637: 9 - Posthumus 965: 7c; 2028: 7c;

2029: 4; 2362: 7c; 2708: 7c; 3155: 4 - Proppe 18: 4 - Pulle 77: 7c; 166: 4 - Pullen 703:

7d; 979: 1; 1323: 7c; 1325: 4; 1327: 6; 1656: 5; 2726: 4; 2739: 5; 3638: 4; 6428: 7d; 6436:

9; 6709: 7d; 6797: 6; 7176: 7c; 7383: 7c; 7429: 4; 8087: 5 - van der Pijl 909: 4. - Raap 35: 1 Rahmat si Boeea 1047: 4/7c; 1259: 7c;4580: 4; 8211: 4 - Rahmat si Toroes, see - praec. - Ramlan 051: 7a - Ramos 1700: 1; 1710: 5 - Rant 15: 1; 17: 7c - Rau 40: 4

Reinwardt 66: 7a - Richards 1104: 1 — Ridley 31: 4; 444: 9; 1111: 4; 1417: 9; 1702: 9;

11525 (Machado): 7c; 11680: 7c; 12340: 7c - Robbins 145: 4; 541: 4; 550: 1 - Robinson - 1651: 7c - Roesil 605: 7c; 717: 7c; 778: 4/7c; 824: 7c; 900: 7c - Romer 358: 4; 623: 4

Roos 6: 7c - Rostados 38: 7c - van Royen 3069: 1; 3109: 9; 3131: 1; 3368: 1; 3485: 1;

1 4757: 7c; 4757a: 7c; 5076: 1; 5287: 4 - RSNB 1220 (Chew et al.): 1; 1418 (id.): -

Rutten-Kooistra 15: 4 - Ruttner 92: 5.

S 17705 (Ashton): 1; 33153 (Chai et al.): 1 - SAN 39356 (Meijer): 7c; 39369 (id.): 7c; 39816

(Madani): 7c; 81304 (Aban & Leopold): 4 - Sands 1881: 7c - Santos 5: 5;4128: 7c;4171:

1; 4175: 7c; 4293: 4; 4508: 4; 4509: 5; 4636: 7c; 4673' 9; 4716: 7c; 4747: 7c; 4850: 1;

4874: 5; 4957: 7c; 4979: 1;4985: 7c; 5011: 4; 5059: 1;5061: 2;5110: 2;5295: 7c;5521:

5/7c; 5592: 5; 5594: 5/7c; 5799: 2;5866: 7c; 6004: 7c;6005: 4;6015a: 1;6096: 7c;6231:

1; 6284: 4; 6287: 7c; 6361: 7c; 6363: 4; 6476: 7c; 6476: 7c; 6513: 1; 6555: 4; 6608: 7c;

6619: 4; 6623: 7c; 6624: 4;6753: 7c;6760: 1;6786: 1;6814: 7c;6861: 7c;6862: 4;6893:

7c; 6984: 4; 6998: 7c; 7072: 5; 7175: 7c; 7429: 4; 7526: 9; 7589: 7c; 7607: 7c; 7632: 7c;

7633: 7c; 7667: 5; 7668: 1; 7669: 4; 7671: 4/7c; 7732: 4; 7747: 4; 7748: 4; 7760: 2; 7776:

5; 7787: 7c; 7844: 2; 7873: 2; 7984: 4; 7985: 7c; 7992: 8; 8028: 8; 8059: 7c; 8067: 7c;

8092: 4; 8102: 7c; 8115: 4/7c; 8136: 9; 8139: 7c; 8150: 4; 8161: 9; 8198: 7c - Sapiin

- 2024: 2 Saunders 99: 4 - Sauveur & Sinke 2518: 6; 2593: 5; 2608: 1 - Saviniere 1660:

7c - Schiffner 1508: 7c; 1540: 1 — Schlenker 29: 7c — Schmutz 1527: 1 — 2080a: 4; 2080b:

7c; 3647: 5; 5000: 7a; 5005: 7c; 5013: 4; 5093: 1; 5265: 5; 5270: 2; 5578: 5; 5579: 7c;

5580: 5; 5584: 4; 6008: 1 - Schodde2153: 6 -Seimund93: 7c; 110: 7c; 316: 7c - SF 363

(Burkill): 4; 573 (id.): 7c; 1275 (id.): 9; 1849 (Alvins): 5;4508 (Nur): 9;4509 (id.): 4;4515 318 BLUMEA VOL. 30, No. 2, 1985

(id.): 7c; 4550 (id.): 7c; 4618 (Burkill): 4;4627 (id.): 7c; 4639 (id.): 7c;4665 (id.): 7c; 4671

(id.): 4; 4672 (id.): 7c; 4751 (Nui): 7c; 4761 (id.): 4; 4779 (id.): 7c; 4781 (id.): 7c; 4784

(id.): 4; 4787 (id.): 7c; 5263 (id.): 7c; 5772 (Native coll.): 4; 6213 (Nur): 7c. 6407 (Burkill):

7c; 6622 (Nur): 3; 6624 (id.): 3; 10153 (Henderson): 4; 10184 (id.): 7c; 13065 (Burkill): 9;

14626 (Boden Kloss): 7c; 17142 (Burkill): 4; 18297 (Henderson): 7c; 19849 (Holttum): 7c;

20215 (Henderson): 7c; 20285 (id.): 4; 21508 (Holttum): 7c; 23752 (Meijer): 1; 24428

(Henderson): 5; 24580 (Holttum): 7c; 24654a (id.): 7c; 24677 (id.): 7c; 24736 (id.): 7c;

25778 (Corner): 7c; 25793 (id.): 9; 25817 (id.): 9; 25821 (id.): 7c; 28530 (id.): 9; 34100

(Nur): 4; 34181 (id.): 4; 35893 (Nauen): 7c; 37331 (Spase): 7c; 37477 (Agric. Off.): 9;

38084 (Nauen): 9; 38190 (id.): 7c; 38192 (id.): 4; 38225 (Henderson): 4; 38858 (Sinclair):

- 4; 40964 (Gilliland): 4; 40967 (id.): 9 - Simpson 8: 7c - Sinke 39: 1 van Slooten 15: 7c;

2028: 9 - Soares 703: (1); 733: (4) - Soedarsono 273: 1; 355: 1 - Soegeng 214: 7c - Sohns

12: 7c; 13: 7c; 14: 4; 29: 7a; 51: 7c; 52: 7c - van Steenis 450: 5;451: 7c; 463: 7c; 820: 7c;

989: 7c; 1012: 1; 3117: 7c; 6452: 5; 6651: 7c; 6655: 7a; 6672: 4; 10583: 9; 11396: 9;

11793: 7c; 18090: 7c; 18091: 4; 18225: 1; 18367: 1; 18368: (2) - Sterly 1658: w - Stevens

et - - al. 712: 7c; 714: 1 Stone 4807: 1 Stresemann 345: 1 - Surbeck 521: 5. - - Taylor 2493a: 7c Teruya 2746: 2 Toroer, see Rahmat si Boeea.

University of San Carlos 807: 1; 873: 7c - UPNG 324 (Tippett): 4; 3219 (Vinas): 4: 3938 (id.):

1; 7564 (Leach): 5.

Vanoverbergh 142: 7c; 811: 5 - Veeartsenijk. Dienst Roeteng 1: 7c; 23: 4; 54: 4 Veemantri

DJ-15: 6; DJ-19: 7c; DJ-20:7c- Veldkamp6971:9;6998:n-Verboom41:4- Verdcourt

5156d: 4 - Verheijen 1529: 7b; 2722: 7c; 3559: 4; 3653: 7b; 4458: 7b; 4460: 4;4474: 1 -

Versteegh 73: 1; 76: 9; 79: 7c; 1162: 7c; 1946: 4 - de Vogel 997: 1; 3036: 1; 3168: 7c;

3410: 1; 3818: 7c;3819: 1.

Walsh 14: 7c; 41: 4; 78: 4; 80: 7c - Weber 1063: 4 - Weinland 192: 7c - Wenzel 1078: 7b;

1349: 7c; 1500: 7c - Whitehead & Flenley JW-112: 5 - Whitford 573: (1) - Widjaja 508: 9

- Wight KD 3031: 5/7c - de Wilde & Vervoort 393: 6; 463: 1 - de Wilde & de Wilde-Duyfjes

12296: 7c; 12303: 1 - Williams 89: 7c; 1177: 5; 1178: 5 - Winckel 58b: 7c; 1029: 7c -

Hans Winkler 420: 1 - Hub. Winkler 2181: 4; 3377: 1 - Wiriadinata 579: 1 - Wisse 670: 7a;

707: 4; 984: 7c - de Wit 278: 4; 4049: 1; 4110: 4; 4112: 7a;4120: 7c;4127: 1;4221: 7c; - 4236: 1;4205: 5; 4286: 7c Wong Yun Siew E13A: 7c.

Yapp 218: 1; 354: 7c.

- van Zanten 1033: 7c; 1055: 1; H-46: 1 Zollinger48: 7b; 192: 4 - Zwaardemaker 5: 1;6: 7c.